Brain Data

Species name
Total CNS volume_1
Total CNS volume_2
Total CNS volume_3
Mantle length_1 (brain size specific)
Mantle length_2 (brain size specific)
Mantle length_3 (brain size specific)
Optic lobe (Wirz, 1959, Table V)
CNS volume mm3 (without optic lobe) "plus grand" (Wirz, 1959)
CNS volume mm3 (without optic lobe) "plus petit" (Wirz, 1959)
CNS volume mm3 (including optic lobe) "plus grand" (Wirz, 1959)
CNS volume mm3 (including optic lobe) "plus petit" (Wirz, 1959)
CNS mantle length "plus grand" (Wirz, 1959)
CNS mantle length "plus petit" (Wirz, 1959)
Brain Data Source
Notes on brain data
Abdopus capricornicus 178.5 118.7 118.2 45 39 38 C22-C22-C22
Abralia veranyi 77.76 70.416 50 42 332 18 16.3 77.76 70.416 50 42 W-W
Abraliopsis morisii 53.946 41.8 42.282 42 32 31 386 11.1 8.7 53.946 42.282 42 31 W-MY-W NY cites MY slightly wrong for the optic lobe, which MY writes as 32 mm^3, not 33 mm^3
Alloteuthis media 184.46 131.56 87.2 70 55 60 360 40.1 28.6 184.46 131.56 70 55 W-W-MY Alloteuthis media is not mentioned in Nixon and Young (2003)
Alloteuthis subulata 74.052 41.14 40 32 384 15.3 8.5 74.052 41.14 40 32 W-W In addition to Wirz data, NY (Table 2.6) has lobe proportions for this species
Amphitretus pelagicus 9.4 32 NY Maddock and Young (1987) does not include ML, but Nixon & Young (2003) does
Ancistroteuthis lichtensteinii 189.224 154.938 160 98 334 43.6 35.7 189.224 154.938 160 98 W-W
Architeuthis dux 1766.6 1570 MY In Nixon and Young (2003) the specimen is listed as Architeuthis sp. even though it is listed as Architeuthis dux by Maddok and Young (1987). TB found that Architeuthis sp. is generally cited as synonymous with A. dux (see e.g. Fonseca et al., 2020, https://doi.org/10.1093/gigascience/giz152) and genetic analyses "strongly suggest" that there is one global species (Winkelmann et al., 2013, https://doi.org/10.1098/rspb.2013.0273).
Argonauta argo 121.66 86.548 34.6 50 40 208 39.5 28.1 121.66 86.548 50 40 W-W-MY
Bathothauma lyromma 46.4 165 MY
Bathypolypus bairdii 193 89 MY Reported for synonym Benthoctopus piscatorum in MY. Muus (2002) determined that Benthoctopus piscatorum (Verrill, 1879) is a junior synonym for Bathypolypus bairdii.
Bathypolypus sponsalis 78.376 51.992 38 45 30 25 94 40.4 26.8 78.376 51.992 45 30 W-W-MY
Bathyteuthis abyssicola 7 30 NY Maddock and Young (1987) merely writes the genus, which is why Nixon and Young (2003) is referenced here
Bolitaena pygmaea 28.8 75 NY In Nixon & Young (2003), the species is reported as Eledonella pygmaea, which in Catalogue of Life and WoRMS is reported as a synonym for Bolitaena pygmaea
Callistoctopus macropus 116.4 MY In Maddock and Young (1987) and Nixon & Young (2003), the species is reported as Octopus macropus, which in Catalogue of Life and WoRMS is reported as a synonym for Callistoctopus macropus
Chiroteuthis veranii 50.2 68 MY
Chtenopteryx sicula 24.2 36 MY
Cirroteuthis muelleri 12 75 MY In Maddock and Young (1987) and Nixon and Young (2003) it is mentioned as Cirroteuthis sp. Cirroteuthis muelleri is the only species in the genus, which is why Cirroteuthis sp.= Cirroteuthis muelleri.
Cirrothauma murrayi 8.2 155 MY
Cranchia scabra 138.8 125 MY
Discoteuthis laciniosa 97.6 51 MY
Egea inermis 53.2 198 MY
Eledone cirrhosa 394.68 187.44 110 66 164 149.5 71 394.68 187.44 110 66 W-W
Eledone moschata 276.336 150.252 59.2 107 66 128 121.2 65.9 276.336 150.252 107 66 W-W-MY
Enteroctopus dofleini 1338.8 MY In Maddock and Young (1987) and Nixon & Young (2003), the species is reported as Octopus dofleini, which is an older name for Enteroctopus dofleini (Catalogue of Life, WoRMS, Jereb et al., 2014)
Galiteuthis glacialis 96.4 297 MY
Gonatus fabricii 101.4 110 MY
Grimalditeuthis bonplandi 27.8 67 MY
Haliphron atlanticus 160.2 MY In Maddock and Young (1987) and Nixon & Young (2003), the species is reported as Alloposus mollis, which in Catalogue of Life and WoRMS is reported as a synonym for Haliphron atlanticus
Hapalochlaena fasciata 54.9 49.7 36 28 C22-C22
Helicocranchia papillata 7.2 MY
Heteroteuthis dispar 42.642 29.394 12.6 20 16 11 314 10.3 7.1 42.642 29.394 20 16 W-W-MY
Histioteuthis miranda 131.6 60 MY
Illex coindetii 447.888 197.532 164 80 272 120.4 53.1 447.888 197.532 164 80 W-W
Illex illecebrosus 198.4 162 MY
Japetella diaphana 5.2 25 NY Maddock and Young (1987) merely writes the genus, which is why Nixon and Young (2003) is referenced here
Joubiniteuthis portieri 7 36 MY
Leachia dislocata 17.8 132 NY Maddock and Young (1987) does not include ML, which is why Nixon and Young (2003) are referenced
Loligo forbesii 283.6 50 MY
Loligo vulgaris 703.984 447.488 245 166 268 191.3 121.6 703.984 447.488 245 166 W-W
Lolliguncula brevis 30.8 75 MY
Lycoteuthis lorigera 189.8 80 MY In Maddock and Young (1987) the species is listed under its synonym Lycoteuthis diadema (see Catalogue of Life and WoRMS) . In Nixon and Young (2003) they write Lycoteuthis lorigera
Macrotritopus defilippi 124.032 76.976 36.4 40 25 172 45.6 28.3 124.032 76.976 40 25 W-W-MY In Maddock and Young (1987), Nixon and Young (2003), and Wirz (1959), the species is reported as Octopus defilippi, which in Catalogue of Life and WoRMS is reported as a synonym for Macrotritopus defilippi
Mastigoteuthis schmidti 28.6 60 MY
Megalocranchia maxima 198.4 335 MY In Maddock and Young (1987) and Nixon & Young (2003) the specimen is listed as Megalocranchia sp. TB have found that the (sub-)species seem to be distinguished primarily based on their geographic distribution (e.g., for the species listed in Jeber & Roper, diagnostic features are shared and each refer to the "generic" features). The data has been added to this species, as this is the most researched
Neorossia caroli 21.4 60 MY
Neoteuthis thielei 18 23 MY In Maddock and Young (1987) Nixon and Young (2003) it is mentioned as Neoteuthis sp. Neoteuthis thielei is the only species in the genus, which is why Neoteuthis sp.= Neoteuthis thielei.
Octopoteuthis danae 139.6 70 MY
Octopus bimaculatus 106.8 MY
Octopus salutii 229.06 184.6 154.44 84 65 160 88.1 59.4 229.06 154.44 84 65 W-MY-W
Octopus vulgaris (all subspecies) 538.572 239.316 171.6 155 80 80 122 242.6 107.8 538.572 239.316 155 80 W-W-MY
Ocythoe tuberculata 264 101.108 93.058 44 42 222 31.4 28.9 101.108 93.058 44 42 MY-W-W
Onychoteuthis banksii 40.11 15.8 39 68 282 10.5 40.11 39 W-MY Weird that the almost double lenght specimen in NY has a brain size much smaller than reported in W
Pickfordiateuthis pulchella 9.2 21 MY
Pteroctopus tetracirrhus 283.528 217.892 21.6 92 70 144 116.2 89.3 283.528 217.892 92 70 W-W-MY
Pterygioteuthis giardi 5.6 23 MY
Pyroteuthis margaritifera 48.888 43.848 19.4 40 32 24 404 9.7 8.7 48.888 43.848 40 32 W-W-MY
Rossia macrosoma 192.808 128.436 77 51 208 62.6 41.7 192.808 128.436 77 51 W-W
Sandalops melancholicus 34.2 112 MY In Maddock and Young (1987) and Nixon and Young (2003) it is mentioned as Sandalops sp. Sandalops melancholicus is the only species in the genus, which is why Sandalops sp.= Sandalops melancholicus
Scaeurgus unicirrhus 138.6 136.25 110.25 62 50 150 54.5 44.1 136.25 110.25 62 50 MY-W-W
Sepia bandensis 312.100 60 M Mantle length of the eight specimens were not measured but estimated to be around 6 cm (pers. comm.).
Sepia elegans 128.7 115.632 55 49 296 32.5 29.2 128.7 115.632 55 49 W-W
Sepia officinalis 433.236 396.2 144.096 185 80 70 216 137.1 45.6 433.236 144.096 185 70 W-MY-W
Sepia orbignyana 245.344 207.196 80 62 274 65.6 55.4 245.344 207.196 80 62 W-W
Sepia plangon 872.5 386.3 455.62 107 72.9 71.1 C23-C23-C23
Sepietta oweniana 72.168 44.268 33 23 272 19.4 11.9 72.168 44.268 33 23 W-W
Sepiola affinis 66.906 30.996 30 19 278 17.7 8.2 66.906 30.996 30 19 W-W
Sepiola robusta 47.104 32.016 24 20 268 12.8 8.7 47.104 32.016 24 20 W-W
Sepiola rondeletii 58.89 36.27 18.2 27 21 23 290 15.1 9.3 58.89 36.27 27 21 W-W-MY
Sepioteuthis lessoniana 582.3848 113 C20 Used Squid 5 in Table S1, since this is the adult
Sepioteuthis sepioidea 261.2 150 MY
Spirula spirula 22.6 21 MY
Taonius pavo 228.4 540 MY
Teuthowenia megalops 260.8 265 MY
Todarodes sagittatus 780.624 631.152 290 235 332 180.7 146.1 780.624 631.152 290 235 W-W In Wirz (1959), the species is reported as Ommatostrephes sagittatus, which in Catalogue of Life and WoRMS is reported as a synonym for Todarodes sagittatus
Todaropsis eblanae 481.954 334.4 148 106 318 115.3 80 481.954 334.4 148 106 W-W
Tremoctopus violaceus 332.008 158.4 24.44 71 15 276 88.3 6.5 332.008 24.44 71 15 W-MY-W The smallest brain data from Wirz is for a male. It's quite a bit smaller than the other two estimates, due to the extreme sexual dimorphism in this species.
Vampyroteuthis infernalis 47 45 MY
Vitreledonella richardi 16.6 66 NY Maddock and Young (1987) merely writes the genus, which is why Nixon and Young (2003) is referenced here

Main Data

Species name
Phylo
Max depth
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Min depth
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Pelagic/Benthic
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Latitude Range
Max latitude
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Min latitude
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Depth category (1 = shallow, 2 = daily mig, 3 = ontogenetic migration, 4 = deep)
Horizontal ontogenetic migration
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Vertical ontogenetic migration
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Vertical daily migration
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Life span max (days)
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Life span min (days)
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Age of sexual maturity max (days)
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Age of sexual maturity min (days)
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Foraging behaviors (summed hunting and feeding behaviors + hunting/feeding strategies)
Hunting and feeding behavior
Preparatory: Ambush/sit-and-wait
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Preparatory: Luring
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Preparatory: Trawl
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Preparatory: Searching
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Preparatory: Follow
other predator or disruption. Scavenging
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Near Contact: Changing
substrate by blowing
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Near Contact: Changing
substrate by digging
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Near Contact: Pursuit
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Near Contact: Stalking
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Near Contact: Visual startle
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Near Contact: Speculative hunting
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Capture Method: Tentacle Strike
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Capture Method: Arm grab
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Capture Method: Trap by web envelop
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Capture Method: Trap by waterflow
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Capture Method: Pull from substrate
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Preparation: Drill/poison (which might immobilize prey)
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Preparation: Remove from shelter/protection
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Preparation: Tactical biting
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Preparation: Pull prey apart
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Preparation: Neutralizing defences
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Hunting/feeding: Strategies
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Defense behaviors
Anti-predator behavior: Types
Avoidance: Staying in shelter
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Avoidance: Manipulating or modifying shelter
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Avoidance: Carry shelter
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Avoidance: Change timing with predation risk
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Avoidance: Choosing/being in an area for activity with low-predation risk
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Avoidance: Take prey home for consumption
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Primary Defense: Burying
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Primary Defense: Camouflage by matching to habitat
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Primary Defense: Camouflage by disruptive pattern
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Primary Defense: Countershading/counter-illumination
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Primary Defense: Active transparency
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Primary Defense: Motion Camouflage or Masquerade when moving
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Primary Defense: Mimicry
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Primary Defense: Schooling
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Primary Defence: Sentinel
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Primary Defence: Look repellent
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Primary Defence: Carry repellent
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Primary Defence: Association with unattractive species
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Secondary Defense: Deimatic/dymantic/startle display
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Secondary Defence: Balloon defence
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Tertiary Defense: Inking
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Ink sac
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Tertiary Defense: Jettisoning of eggs
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Tertiary Defense: Flight
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Tertiary Defense: Jumping out of water
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Tertiary Defense: Crawling up on land
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Tertiary Defense: Autotomy
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Anti-predator behavior: Strategic
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Diet breadth: Species richness
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Data from midden
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Data from stomach
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Basic prey species
Number of predator species
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Basic predator groups
Sociality type
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Sociality type (binary)
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Combined cognition
Cognition Sum
Habituation
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Sensitization
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Classical conditioning
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Instrumental/operant conditioning
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Avoidance learning
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Conditional discrimination
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Episodic-like memory
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Reversal learning
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Spatial learning
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Visual learning
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Chemical modality
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Mechanoreceptive/tactile modality
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Self-recognition
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Short-term memory
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Long-term memory
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Imprinting
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Play
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Tool use
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Contra-freeloading
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Number sense/quantity recognition
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Problem solving
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Mazes
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Deception
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Bilateral signal control
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Social sum
Between species association
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Sociality dynamics: Individual recognition
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Non-agonistic communication w. conspecifics (non-reproductive context)
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Agonistic communication w. conspecifics (non-reproductive context)
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Number of displays
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Aggregating for mating/spawning
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Positive behaviour (female to male)
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Negative behaviour (female to male)
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Positive behaviour (male to female)
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Negative behaviour (male to female)
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Competition between males
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Competition between females
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Cryptic female choice
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Mate guarding
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Sneaker male
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Female mimicry
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Alternating between consort and sneaker strategy depending on context
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Egg care (carrying eggs)
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Egg care (eggs attached to substrate)
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Research effort (Web of Science citations, species name and alternative name in topic 1866-2020)
Articles read
Total CNS volume_1
Total CNS volume_2
Total CNS volume_3
Mantle length_1 (brain size specific)
Mantle length_2 (brain size specific)
Mantle length_3 (brain size specific)
Brain Data Source
Notes on brain data
Abdopus capricornicus Abdopus aculeatus 10 "It has been observed to forage over intertidal coral bedrock and coral
rubble flats during daylight low tides. " (Norman & Finn, 2001)		 (Norman & Finn, 2001) 0 "It has been observed to forage over intertidal coral bedrock and coral
rubble flats during daylight low tides. " (Norman & Finn, 2001)		 (Norman & Finn, 2001) 2 "It has been observed to forage over intertidal coral bedrock and coral
rubble flats during daylight low tides. " (Norman & Finn, 2001)		 Norman & Finn, 2001
Scata, 2022		 0.5 -23 "Known only from the intertidal reefs surrounding coral
islands in the Capricorn Bunker island group at the southern
end of the Great Barrier Reef, Australia." (Norman & Finn 2001)		 (Norman & Finn 2001; Jereb et al., 2014) -23.5 "Australia, known only from Tryon Island, Great Barrier Reef. " (Jereb et al., 2014) (Norman & Finn 2001; Jereb et al., 2014) 1 365.3 TB: I wrote Wen-Sung Chung who supervised the recent doctoral thesis on this species whether they had good guesses about the life span and/or age of sexual maturity. Unfortunately, no one seems to know much about the life-history of A. capricornicus (see Wen-Sung's reply in quotes below). An alternative lead would be Christine Huffard who's done detailed work on a similar species, A. aculeatus. "Dear Theiss,

Thanks for your mail.

Re the life spaNAge question, honestly, we/I have no clear clue for how long they can live. Firstly, this species lays small eggs in terms of difficulties to keep planktonic larvae alive and no success on raising them up yet. Second, we have collected this species over the past 5 years at the same place during the period of the annual greatest spring tide. Although we also had tried different seasons to hunt them, given the experienced eyes, we barely saw them due to their bloody good camouflage under high water level and big swell conditions. I would guess their life span is about 1 yr as the maturity of most animals we collected was within a similar stage (ready for mating). This is the only clue which we have, sorry.

Best

Wensung"

 pers. comm. 365.3 TB: I wrote Wen-Sung Chung who supervised the recent doctoral thesis on this species whether they had good guesses about the life span and/or age of sexual maturity. Unfortunately, no one seems to know much about the life-history of A. capricornicus (see Wen-Sung's reply in quotes below). An alternative lead would be Christine Huffard who's done detailed work on a similar species, A. aculeatus. "Dear Theiss,

Thanks for your mail.

Re the life spaNAge question, honestly, we/I have no clear clue for how long they can live. Firstly, this species lays small eggs in terms of difficulties to keep planktonic larvae alive and no success on raising them up yet. Second, we have collected this species over the past 5 years at the same place during the period of the annual greatest spring tide. Although we also had tried different seasons to hunt them, given the experienced eyes, we barely saw them due to their bloody good camouflage under high water level and big swell conditions. I would guess their life span is about 1 yr as the maturity of most animals we collected was within a similar stage (ready for mating). This is the only clue which we have, sorry.

Best

Wensung"

 pers. comm. 5 5 1 "Moving rock" camouflage (Scata, 2022, Table 4.4) NB: purpose (defense, foraging, etc.) not mentioned specifically Scata 2022 1 "It has been observed to forage over intertidal coral bedrock and coral rubble flats during daylight low tides." (Norman & Finn 2001)" Norman and Finn 2001
Chung et al. 2022		 1 digging/burying (Scata, 2022, Table 4.4) NB: purpose (defense, foraging, etc.) not mentioned specifically
inking (Scata, 2022, Table 4.4) NB: purpose (defense, foraging, etc.) not mentioned specifically		 Scata 2022 1 chasing, charging (Scata, 2022, Table 4.4) NB: purpose (defense, foraging, etc.) not mentioned specifically Scata 2022 1 webover (Scata, 2022, Table 4.4) NB: purpose (defense, foraging, etc.) not mentioned specifically Scata 2022 6 6 1 "Moving rock" camouflage (Scata, 2022, Table 4.4) NB: purpose (defense, foraging, etc.) not mentioned specifically Scata, 2022 1 digging/burying (Scata, 2022, Table 4.4) NB: purpose (defense, foraging, etc.) not mentioned specifically Scata, 2022 1 "sophisticated camouflage to look like seaweed—hence, its other common name, the algal octopus" (Chung et al. 2022) Chung et al. 2022 1 "Flushing", i.e. jetting water at object or another animal (Scata, 2022, Table 4.4) NB: purpose (defense, foraging, etc.) not mentioned specifically Scata, 2022 1 inking (Scata, 2022, Table 4.4) NB: purpose (defense, foraging, etc.) not mentioned specifically Scata, 2022 1 Present in the genus (Jereb et al., 2014) (Jereb et al., 2014) 1 escape (Scata, 2022, Table 4.4) NB: purpose (defense, foraging, etc.) not mentioned specifically Scata, 2022 0 0 2 "[A. capricornicus is] found in aggregations with a high degree of conspecific interactions…Unlike most octopus species which are solitary through almost their entire lifespan, the Abdopus mating system has much in common with that of gregarious cephalopods such as loliginid squid (Hanlon & Messenger, 2018)." (Scata, 2022: 190, 276) 0 "[A. capricornicus is] found in aggregations with a high degree of conspecific interactions…Unlike most octopus species which are solitary through almost their entire lifespan, the Abdopus mating system has much in
common with that of gregarious cephalopods such as loliginid squid (Hanlon & Messenger, 2018)." (Scata, 2022: 190, 276)		 3 0 3 0 "A. capricornicus did not engage in less interactions with the familiar conspecific compared to those in unfamiliar trials. In addition, latencies of first approach and of first physical contact were not longer in familiar trials as expected if animals are capable of recognizing conspecifics." (Scata, 2022: 267) Scata, Gabriella. “Brain Structure, Body Patterns and Social Interactions in the Octopus Abdopus Capricornicus.” PhD Thesis, The University of Queensland, 2022. https://doi.org/10.14264/053d1ea. 1 "Males show highly contrasting color patterns (e.g., a white body with dark longitudinal stripe(s) from mantle to arm crown) along with frequent physical interactions (e.g., arm touching) with potential mates. The dominant male often shows strong mate guarding to
keep his mate within arm-reaching range and frequently mates
over successive days. 99" (Chung et al., 2021)		 (Chung et al., 2021) 1 "Mating interactions of A. aculeatus have previ-ously been recorded in its natural habitats (Indonesia) such as male competitions, mate guarding, and other tactics such assneaker male copulations, similar to those in socially aggre-gating cuttlefish or squid.39,53,55,110,111We have documented similar social interactions in A. capricornicus.99 Males show highly contrasting color patterns (e.g., a white body with dark longitudinal stripe(s) from mantle to arm crown) along withfrequent physical interactions (e.g., arm touching) with potential mates. The dominant male often shows strong mate guarding to keep his mate within arm-reaching range and frequently matesover successive days.99These unique paired-pond behaviors could ensure reproductive success, and the effects of social or-ganization may therefore drive differentiation of cognition devel-opment similar to the examples seen in other invertebrates suchas solitary versus eusocial insects" (Chung et al. 2022). [TB: Note that Jennifer reviewed the thesis which this claim is based on and was not at all impressed with the quality of the research] (Chung et al. 2022) 1 "Males show highly contrasting color patterns (e.g., a white body with dark longitudinal stripe(s) from mantle to arm crown) along with frequent physical interactions (e.g., arm touching) with potential mates. The dominant male often shows strong mate guarding to
keep his mate within arm-reaching range and frequently mates
over successive days. 99" (Chung et al., 2021)		 (Chung et al., 2021) 1 2 178.5 118.7 118.2 45 39 38 C22
Abralia veranyi Abralia veranyi 732 588-732 m (Vecchione & Roper, 1991, p. 436) (Vafidis et al 2008 0 "Depths of capture have ranged between the surface and 550 m." (Cairns, 1976) Cairns, 1976 1 "At bottom in bathyal and in midwater above slopes, occasionally to surface (night time); absent in the open ocean far from the slopes and slope waters." (Jereb & Roper, 2010) Jereb & Roper, 2010 44 38 MessiNA"Mediterranean Sea, off Messina, Italy; Atlantic Ocean. Tropical and subtropical western and eastern Atlantic Ocean, from northeastern United States to the Gulf of Mexico and Suriname; from Mediterranean Sea to Madeira." (Jereb & Roper, 2010) (Jereb & Roper, 2010) -6 Congo River "A. veranyi has been reported from the Mediterranean and both sides of the Atlantic from the Bay of Funchal, Madeira, to off the Congo River in the Eastern Atlantic, and in the Western Atlantic in the Gulf of Mexico, off Cuba and off Key West (Voss, 1956a). " (Cairns 1976) (Cairns 1976) 2 1 " Our study has shown
that the cephalopods A. veranyi feed between
dusk and dawn in the epipelagic zone (0–200 m depth), pointing to
the possibility that these species play a key role in the oceanic food
web (Clarke, 1996; Ariza, 2015) and in the sequestration of carbon.
We found that individuals of both species exhibited greater stomach
fullness during nighttime hours compared with daytime hours, and
this observation is consistent with vertical migration to the surface
at night to feed" (Guerra-Marrero et al 2020)		 (Guerra-Marrero et al 2020) (Jereb & Roper, 2010) NA 0 1 "mediumsized individuals and adults of A. veranyi preyed on fish at any time of the day, but on cephalopods only during daylight hours" (Guerra-Marrero et al. 2020) Guerra-Marrero et al. 2020 2 2 1 “The ocular photophore flashes had a broader bandwidth and a shorter wavelength emission maximum than that of the ventral mantle at 11 to 12 °C…Their broad bandwidth emission is not a good match of downwelling daylight and, as we observed, is probably used more often in a defensive or "startle" mode” (Herring et al. 1992) Herring et al. 1992 1 Reported attracted to light bulb underwater and "assumed a type of distinctive "upward V curl" posture (Moynihan, 1975) in which all or most of the arms on both sides of the body were held upward and somewhat backward forming a V (Figure 2C). This posture and its associated color arrangements are thought to be an indication of alarm, possibly allowing the squid to thwart predators by mimicing or hiding in drifting Sargassum weed or gorgonians" (Hanlon et al. 1979) TB: this posture is described as "flamboyant" in Hanlon & Messenger, 2018, p. 126 Hanlon & Messenger, 2018, p. 126 6 "Abralia veranyi feed mainly on zooplankton, particularly on copepods, mysids and the early growth stages of decapod crustaceans. " (Guerra-Marrero et al 2020) Guerra-Marrero et al 2020
Vafidis et al. 2009		 0 1 Vafidis et al. 2009 3 6 made
up about 15% of cephalopod species found in stomachs of demersal sharks
Scyliorhinus canicula and Squalus blainville caught in Aegean Sea (Kousteni et
al. 2018)		 (Kousteni et
al. 2018) (Ozturk
et al. 2007) (Konan
et al. 2018) (de
Gurjao et al. 2004) (Staudinger
et al. 2013) 		3 0 not gregarious (inferred from photo and video material) 1 0 1 0 "As we observed, Abraliopsis morisii and A. veranyi are
distributed throughout the mesopelagic layer during the daytime
and ascend at night to the epipelagic region to feed (the two species
feed at different depth levels and this may reduce interspecific
competition)." (Guerra-Marrero et al 2020)		 Guerra-Marrero et al 2020 1 "A. veranyi, a small mesopelagic squid, was an occasional prey across predators and regions; however, the occurrence of 53 individuals in a single yellowfin tuNAstomach during fall of 2007, and additional incidences during late summer and early fall during other years indicates this species may be aggregating in large numbers, most likely to spawn, in waters off of New England." (Staudinger et al. 2012) (Staudinger et al. 2012) 36 19 77.76 70.416 50 42 W-W
Abraliopsis morisii Abraliopsis morisii 650 "The typical presence of A. morisii, O. sicula, T. megalops, and B. beanii in the shallow night catch but not the shallow day catch (Table 7) is suggestive of diel vertical migration in these species. This result is consistent with previous reports about A. morisii (as A. pfefferi) in closing nets around Bermuda where a 610–650 m daytime and 50–100 m nighttime depth was recorded by Roper and Young (1975)." (Shea et al. 2017) (Shea et al. 2017) 50 "The typical presence of A. morisii, O. sicula, T. megalops, and B. beanii in the shallow night catch but not the shallow day catch (Table 7) is suggestive of diel vertical migration in these species. This result is consistent with previous reports about A. morisii (as A. pfefferi) in closing nets around Bermuda where a 610–650 m daytime and 50–100 m nighttime depth was recorded by Roper and Young (1975)." (Shea et al. 2017) (Shea et al. 2017) 1 "Oceanic, mesopelagic " (Gomes-Pereira et al., 2016) Brandt 1983
Gomes-Pereira et al., 2016		 51 51 "l”51’ N. 0-31’ E. Atlantic Ocean " (Glaubrecht and Salcedo-Vargas 2000) (Glaubrecht and Salcedo-Vargas 2000) 0 Natal is at equator "A. pfefferi is known from the Mediterranean (Joubin, 1896), the northeastern Atlantic (Joubin, J920, 1924) and
the lndian Ocean SSE of Natal (Voss, 1967a). Three adult specimens were taken from the Straits in Northern Yucatan and Transitional waters. The other specimen was captured north of Little Bahama Bank. This is the first record of A. pfefferi from the Western Atlantic." (Cairns 1976)		 (Cairns 1976) 2 1 “The typical presence of A. morisii, O. sicula, T. megalops, and B. beanii in the shallow night catch but not the shallow day catch (Table 7) is suggestive of diel vertical migration in these species. This result is consistent with previous reports about A. morisii (as A. pfefferi) in closing nets around Bermuda where a 610–650 m daytime and 50–100 m nighttime depth was recorded by Roper and Young (1975)." (Shea et al. 2017) (Shea et al. 2017) 154 "In our samples, minimum age (86 d) was observed both in a maturing male of 22 mm ML and a maturing female of 24 mm ML. Maximum age was revealed in a mature male of 25 mm ML (127 d) and in a mature female of 33 mm ML (154 d)" Arkhipkin (1996), p. 327 Arkhipkin (1996), p. 327 86 "In our samples, minimum age (86 d) was observed both in a maturing male of 22 mm ML and a maturing female of 24 mm ML. Maximum age was revealed in a mature male of 25 mm ML (127 d) and in a mature female of 33 mm ML (154 d)" Arkhipkin (1996), p. 327 Arkhipkin (1996), p. 327 127 Min. observed: 100 days, max. observed: 127 days (Arkhipkin (1996), p. 328-329) (Arkhipkin (1996), p. 328-329) 100 Min. observed: 100 days, max. observed: 127 days (Arkhipkin (1996), p. 328-329) (Arkhipkin (1996), p. 328-329) NA 0 1 1 1 species unspecified "Abraliopsis living in a shipboard aquarium was observed to match the overhead illumination with a downward glow, so that to the human observer the squid ‘disappeared’ (R.E. Young and Roper 1977, R.E. Young et al. 1979a). The values of maximum illumination intensity obtained for this squid indicated that the limits for concealment from predators determine the upper limits which it can inhabit during the day (R.E. Young et al. 1980, R.E. Young and Mencher 1980)." (Nixon & Young 2003) Nixon & Young 2003 7 Bear Seamount "feeding primarily on copepods, euphausiids, other small invertebrates, and to a lesser extent small fishes and cephalopods (Passarella & Hopkins 1991)…For the early to mid-life stages represented in our evaluations,these squids appear to be primarily tracking and consuming lower trophic level prey (e.g., zooplankton) as they migrate vertically in the water column" (Staudinger et al. 2019) & " Abraliopsis morisii feed mainly on zooplankton,
particularly on copepods, mysids and the early growth stages of
decapod crustaceans." (Guerra-Marrero et al 2020)		 (Staudinger et al. 2019) & (Guerra-Marrero et al 2020) 0 1 (Guerra-Marrero et al 2020) 5 1 "Risso’s dolphin, Grampus griseus (Cuvier, 1812)" (Blanco et al., 2006) Blanco et al., 2006 1 0 not gregarious (inferred from photo and video material) NA 0 NA 0 "As we observed, Abraliopsis morisii and A. veranyi are
distributed throughout the mesopelagic layer during the daytime
and ascend at night to the epipelagic region to feed (the two species
feed at different depth levels and this may reduce interspecific
competition)." 		(Guerra-Marrero et al 2020) 16 11 53.946 42.8 42.282 42 32 31 W-NY-W
Alloteuthis media Alloteuthis media 674 Caught in depths of 20–674 m with mean depth of 97 m (Silva et al., 2011) (Jereb & Roper, 2010; Arkhipkin et al., 2015; Quetglas et al., 2000; Sifner et al., 2005; Silva et al., 2011; Salman et al. 2002; Ruby and Knudsen 1972; Gestal et al., 1999; Arkhipkin and Nekludova, 1993; Rosas-Luis & Sanchez, 2014; Jereb et al., 2015; Katsanevakis et al 2008; Roper & Young, 1975; Vafidis et al 2008; Pierce et al., 2010) 0 "Alloteuthis media is found from the surface to about
500 m depth (Pierce et al. 2010). This vertical
distribution supports their importance as a food
resource in the trophic web, and as predators they
could prey on fish, molluscs and crustaceans, similarly to other squid (Rasero et al. 1996; Nyegaard
2001; Rosas-Luis et al. 2014a, 2014b), but data on
their feeding habits are absent (Jereb & Roper 2010)" (Rosas-Luis & Sanchez, 2014)		 (Jereb & Roper, 2010; Arkhipkin et al., 2015; Quetglas et al., 2000; Sifner et al., 2005; Silva et al., 2011; Salman et al. 2002; Ruby and Knudsen 1972; Gestal et al., 1999; Arkhipkin and Nekludova, 1993; Rosas-Luis & Sanchez, 2014; Jereb et al., 2015; Katsanevakis et al 2008; Roper & Young, 1975; Vafidis et al 2008; Pierce et al., 2010) 2 **edge case** "occurs on sandy and muddy grounds and preferentially inhabits coastal and shelf waters (from the surface to 200 m), even though it has been recorded down to about 500 m depth." (Jereb & Roper, 2010) Jereb & Roper, 2010
Arkhipkin and Nekludova, 1993
Katsanevakis et al 2008		 39 60 "widely distributed in the eastern Atlantic, from the north-western coast of Africa (21°N) to the North Sea (60°N) (Guerra 1992). It is most common south of 50°N, in the Bay of Biscay, English Channel and throughout the Mediterranean." (Hastie et al. 2009) (Hastie et al. 2009) 21 "widely distributed in the eastern Atlantic, from the north-western coast of Africa (21°N) to the North Sea (60°N) (Guerra 1992). It is most common south of 50°N, in the Bay of Biscay, English Channel and throughout the Mediterranean." (Hastie et al. 2009) (Hastie et al. 2009) 1 1 "It migrates towards the coast in March for breeding, leaving again in August. In December they are found at depths of about 100m. " (Ruby and Knudsen 1972) (Ruby and Knudsen 1972; Jereb & Roper, 2010; Jaeckel, 1937; Pierce et al., 2010; Jereb et al., 2015; Katsanevakis et al 2008) 1 "Adults of A. media migrate to shallow water, spawning at depths of 10 – 100 m on sand, seagrass meadows, etc.
from March to October in the western Mediterranean (Mangold‐Wirz, 1963) and year‐round in the central and eastern region (Lo Bianco, 1909; Naef, 1923; Laptikhovsky et al., 2002; Lefkaditou, 2006)." (Pierce et al., 2010)		 (Pierce et al., 2010; Jereb et al., 2015) 547.9 "Based on length frequency analyses, the longevity of A. media had been estimated to be
ca. 1 year for males and 18 months for females, with a monthly growth rate decreasing
from 7–8 mm in the first summer of life to 2–5 mm during the second year (Mangold-Wirz, 1963a; Zuev and Nesis, 1971; Auteri et al., 1987). However, recent direct age de-
termination of A. media in the northwestern Aegean Sea, based on enumeration of daily
increments in statoliths, has shown that the lifespan of females reaches up to 11
months, whereas the males can reach 9 months of age (Alidromiti et al., 2009). " (Jereb et al., 2015)		 (Jereb & Roper, 2010; Pierce et al., 2010; Katsanevakis et al 2008; Arkhipkin and Nekludova, 1993; Arkhipkin et al., 2015; Jereb et al., 2015; Laptikhovsky et al., 2002; Arkhipkin et al., 2015) 182.6 "Alloteuthis spp. have a maximum age of around 12 months (Rodhouse et al., 1988; Moreno et al., 2007). The lifecycle lasts between 6 and 12 months;" (Arkhipkin et al., 2015) (Jereb & Roper, 2010; Pierce et al., 2010; Katsanevakis et al 2008; Arkhipkin and Nekludova, 1993; Arkhipkin et al., 2015; Jereb et al., 2015; Laptikhovsky et al., 2002; Arkhipkin et al., 2015) NA 0 NA 0 18 Table 1 from (Rosas-Luis & Sanchez, 2015) & "The diet of A. media consists of larvae and juveniles of 
fish, copepods, and euphausiids (Zuev and Nesis, 1971). " (Pierce et al., 2010)		 (Rosas-Luis & Sanchez, 2015) & (Jereb & Roper, 2010) & (Pierce et al., 2010) 0 1 (Rosas-Luis & Sanchez, 2015) 5 15 found in stomachs of chondrichthyan Raja clavata southern Adriatic Sea (Bello 1997) Bello 1997
Quetglas et al. (1998)
Sifner & Vrgoc (2009)
Castriota et al. 2015
Velasco et al., 2001
Pierce et al., 2010
Carpentieri et al., 2007
Jereb et al., 2015		 4 0 not gregarious (inferred from photo and video material) NA 0 NA 35 25 184.46 131.56 70 55 W-W
Alloteuthis subulata Alloteuthis subulata 500 "occurs from coastal shallow waters (less than 50 m) down to about 500 m, even though captures below 300 m are sporadic." (Jereb & Roper, 2010) (Jereb & Roper, 2010; Sifner et al., 2005; Silva et al., 2011; Oesterwind et al 2010; Garcia-Mayoral et al., 2020; Lefkaditou et al., 2012; Mathger and Denton, 2001; Gestal et al., 1999; De Heij and Baayen, 1999; Cornwell et al., 1997; Morris et al., 1993; Arkhipkin and Nekludova, 1993; Arkhipkin and Nekludova, 1993; Lipinski, 1985; Llewellyn, 1984; Rodhouse et al., 1988; Adam, 1937; Jereb et al., 2015) 8 "pelagic […] at depths of 8 to 60 m. (trawl)" (Rodhouse et al., 1988) (Jereb & Roper, 2010; Sifner et al., 2005; Silva et al., 2011; Oesterwind et al 2010; Garcia-Mayoral et al., 2020; Lefkaditou et al., 2012; Mathger and Denton, 2001; Gestal et al., 1999; De Heij and Baayen, 1999; Cornwell et al., 1997; Morris et al., 1993; Arkhipkin and Nekludova, 1993; Arkhipkin and Nekludova, 1993; Lipinski, 1985; Llewellyn, 1984; Rodhouse et al., 1988; Adam, 1937; Jereb et al., 2015) 2 **edge case** ""a near-bottom species that lives in shelf waters, particularly in the North Sea (Grimpe, 1925; Steimer, 1993), Kattegat, and western Baltic Sea (e.g. Jaeckel, 1937; Herrmann et al., 2001; Hornbörg, 2005). In UK waters, the species is abundant in the English Channel (Rodhouse et al., 1988) and the Irish Sea (Nyegaard, 2001)." (Jereb et al., 2015) Jereb et al., 2015 40 60 "from approximately 60°N to 20°S:" (Jereb & Roper, 2010) [Alex: The southern distribution looks to be a mistake. On the map, the southest point is 20 N, which fits better with other sources] (Jereb & Roper, 2010) 20 “were collected from the continental shelf off the west Sahara (20°57' to 22°54'N 16°57' to 17°32'W) ” (trawl) (Arkhipkin and Nekludova, 1993) (Arkhipkin and Nekludova, 1993) 1 1 "In some parts of its range, A. subulata is thought to be migratory. For example, in the North Sea, juveniles are thought to leave the area at an age of about 3 mo in November and return the following spring at a length of about 5 cm ML. In this area, males and females move inshore in early summer but are absent during winter (Yau 1994). However, in other areas (e.g., English Channel), this species can be found year-round (Rodhouse et al. 1988)." (Hastie et al. 2009) (Jereb & Roper, 2010; Hastie et al. 2009; De Heij and Baayen, 1999; Jaeckel, 1937; Jereb et al., 2015) 1 "Another opinion is that in response to cooling during late autumn/winter, the juvenile A. subulata migrate from the spawning grounds in the southeastern parts of the North Sea to the deeper, in winter relatively warmer, waters in the central parts (De Heij and Baayen, 2005). In spring the young adults return to the warming shallow waters from the Danish to the Belgian coast and southeastern British coast for spawning (De Heij and Baayen, 1999), which is supported by our data (Fig. 1). During summer, a large number of mature and large A. subulata was encountered in the southeastern parts of the North Sea, whereas in the central and northern parts this species was not common. In winter, A. subulata was caught in large numbers in the central and northern parts, suggesting that it lives permanently in the North Sea with spawning grounds in the southeast in summer and feeding grounds in the central parts in winter. " (Oetserworld et a 2010) (Oetserworld et a 2010; De Heij and Baayen, 1999; Jaeckel, 1937) 730.5 “Maximum length of life for squid from this population is ca 1 yr. […] The disappearance of all sizes of males, apart from recent recruits, in November indicates that the maximum life span is ca 1 yr […] This is consistent with the observation that Alloteuthis subulata lives for 1 to 2 yr (Roper et al. 1984)” (Rodhouse et al., 1988) (Arkhipkin et al., 2015; Arkhipkin and Nekludova, 1993; Jereb et al., 2015; Arkhipkin and Nekludova, 1993; Jereb & Roper, 2010; Arkhipkin and Nekludova, 1993; Pierce et al., 2010; Rodhouse et al., 1988) 172 “In A. subulata the oldest mature male […] was 231 d old, while the oldest mature female […] was 172 d. “ (Arkhipkin and Nekludova, 1993) (Arkhipkin et al., 2015; Arkhipkin and Nekludova, 1993; Jereb et al., 2015; Arkhipkin and Nekludova, 1993; Jereb & Roper, 2010; Arkhipkin and Nekludova, 1993; Pierce et al., 2010; Rodhouse et al., 1988) 180 “Both species matured over a wide range of sizes and ages (from 120 to 180 d) “ (Arkhipkin and Nekludova, 1993) (Arkhipkin and Nekludova, 1993) 90 “Sex of squid could be determined after an age of 90 d “ (Arkhipkin and Nekludova, 1993) (Arkhipkin and Nekludova, 1993) 2 2 1 "Nyegaard (2001) showed that, although A. subulata was associated with the distribution of its main prey species, the squid at stations with high prey abundance did not seem to have been more frequently engaged in feeding activity than those at other stations (based on stomach fullness). This could indicate that A. subulata feeds in the pelagic zone rather than close to the bottom. Indeed both sandeel and sprat, which are important prey of Alloteuthis, undertake vertical migrations and were found in higher abundances in the pelagic zone than near the bottom during the day in the North Sea (Pedersen 1999)." (Hastie et al. 2009) Hastie et al 2009 1 "Food capture was effected by the squid darting forward in a straight line from as far as about 20 cm from the prey;" (LLewellyn, 1984) Llewellyn 1984 4 4 1 "All Dark [components] is seen in squids that are startled or alarmed. "(Cornwell et al., 1997) Cornwell et al., 1997
Mathger, 2003		 1 "The reflections from the iridophores are often inconspicuous, or even absent. If the stripes are fully reflective, however, they produce very clear reflective patterns that contribute substantially to the overall appearance of the animal. The ‘red’ stripes, for example, are often visible when the animal is threatened (e.g. by a hand net) or during intraspecific encounters." (Mathger et al., 2004) Mathger et al., 2004
Lima et al., 2003
Mathger and Denton, 2001
Mathger and Denton, 2001		 1 The parts of the squid that are probably most conspicuous are the £uorescent ‘eyespots’ and the internal organs and it appears likely that using the chromatophores to produce a counter- shading e¡ect is an e¡ective way to conceal these parts of the body. " (Mathger, 2003) Mathger, 2003
Mathger and Denton, 2001
Hanlon & Messenger, 2018, p. 108
Mathger, 2003		 1 “escape reactions, probably including ink-rejection” (Messenger, 1979) Messenger, 1979 1 "Apart from the eyes and the inksac, the body of a squid is very transparent." (Mathger and Denton, 2001) (Mathger and Denton, 2001; Mathger, 2003) 9 "A. subulata show an ontogenetic shift in feeding habits, from a diet dominated by small crustaceans to a mainly piscivorous diet" (Arkhipkin et al., 2015) Arkhipkin et al., 2015
Abbott et al., 1988
Hastie et al., 2009
Jereb et al 2015		 0 0 2 30 "Risso’s dolphin, Grampus griseus (Cuvier, 1812)" (Blanco et al., 2006) [the dolphin was stranded in the western Mediterranean]. "Eledone cirrhosa was the most important component by weight in the stomach contents of an individual stranded on the south coast of England (e.g. Clarke & Pascoe, 1985). In Scottish waters, it is thought to almost exclusively consume a single prey species, E. cirrhosa (Pierce et al., 2007)." (MacLeod et al. 2014); Important prey of Risso's Dolphin (Grampus griseus) in Faroe Islands in April (Bloch et al. 2012) Bloch et al. 2012
Velasco et al., 2001
Xavier et al., 2018
Battaglia et al. 2013
Quigley & Flannery 2014
Villanueva & Norman, 2008
Pierce et al., 2010
Jereb et al., 2015		 4 3 "These squid are schooling animals that need to signal changes in relative position to neighbouring squid to maintain the coherence of the school." Mathger & Denton 2001 (Mathger and Denton, 2001) 1 "These squid are schooling animals that need to signal changes in relative position to neighbouring squid to maintain the coherence of the school." Mathger & Denton 2001 (Mathger and Denton, 2001) 5 1 1 "A very interesting observation was that many squid expanded the chromatophores of the ‘eyespot’ opposite that facing the brightest light (see Figure 4). This, at ¢rst, appears to be the wrong way round. When looking at these squid closely, however, it becomes obvious that the dorsal arches of the eyes are slightly elevated with respect to the rest of the head, so that a predator looking at a squid from an oblique angle would only see the fluorescent layers of the ‘eyespot’ on the opposite side to that facing the brightest light. This would explain why most squid only expand the chromatophores on that side." (Mathger, 2003) Mathger 2003 4 1 “Both sexes appear early in summer for spawning in the southern North Sea. Some adults migrate to coastal waters for egg deposition in September, ” (De Heij and Baayen, 1999) & “There were apparently 3 spawning groups of females in the year sampled. […] which spawned in the spring, summer and autumn.” (Rodhouse et al., 1988) (De Heij and Baayen, 1999) 1 "Several components are used in sexual signalling and are sex-specific. Males may show Accentuated testis, highlighting the white gonad in a translucent body. Females may show a Lateral blush to males. Males show Dorsal mantle splotches when interacting with other males during. Tentacular stripe and spots (not shown) may also be used during agonistic displays (Hanlon, 1982; Lipinski, 1985) " (Cornwell et al. 1997) (Cornwell et al. 1997) 1 "When a single male courted a single female, however, a different pattern [than the 'domination display' of regular dark patches] was observed as in Fig. 7C. Both displays were always supplemented by irregular stitchwork pattern on the fins (Fig. 7A). The courted females never showed patterning, but remained 'all clear' (see also Hanlon, 1982)." (Lipinski, 1985) (Lipinski, 1985) 1 "Several components are used in sexual signalling and are sex-specific. Males may show Accentuated testis, highlighting the white gonad in a translucent body. Females may show a Lateral blush to males. Males show Dorsal mantle splotches when interacting with other males during. Tentacular stripe and spots (not shown) may also be used during agonistic displays (Hanlon, 1982; Lipinski, 1985) " (Cornwell et al. 1997) (Cornwell et al. 1997) 104 80 74.052 41.14 40 32 W-W In addition to Wirz data, NY (Table 2.6) has lobe proportions for this species
Amphitretus pelagicus Amphitretus pelagicus 2000 "Depth range from 100 to 2 000 m" (Jereb et al., 2014) (Jereb et al., 2014; Norman and Reid 2000 – Book, p52) 100 "Depth range from 100 to 2 000 m" (Jereb et al., 2014) (Jereb et al., 2014; Norman and Reid 2000 – Book, p52) 1 "This midwater species lives at depths between the epipelagic and bathypelagic zones" (Jereb et al., 2014) Jereb et al., 2014 72 42 42 N to 30 S (Jereb et al., 2014) (Jereb et al., 2014) -30 42 N to 30 S (Jereb et al., 2014) (Jereb et al., 2014) 4 1 "Juveniles appear to occur in shallower water. " (Jereb et al., 2014) (Jereb et al., 2014) 1 1 1 "The telescopic eyes are always oriented upwards, presumably used to detect the silhouettes of prey from below" (Jereb et al., 2014) Jereb et al 2014 2 1 1 "The octopus minimises its own silhouette by being mostly transparent and by maintaining both eyes and the digestive gland (liver) in a vertical orientation." (Jereb et al., 2014) Jereb et al., 2014 1 Present in the family (Jereb et al., 2014) (Jereb et al., 2014) 0 0 0 not gregarious (inferred from photo and video material) 1 0 1 1 "It is likely that females brood the eggs within their arm crown as has been reported for Vitreledonella." (Jereb et al., 2014) Jereb et al., 2014 0 "It is likely that females brood the eggs within their arm crown as has been reported for Vitreledonella." (Jereb et al., 2014) Jereb et al., 2014 7 2 9.4 32 NY
Ancistroteuthis lichtensteinii Ancistroteuthis lichtensteinii 1271 "One female (129 mm ML) was captured at a depth of 1271 m. " (Villanueva 1992) Jereb & Roper, 2010 0 "800–0m" (Bolstad 2010) Bolstad 2010 1 "an epipelagic, mesopelagic to benthi-bathyal species that occurs in open warm-temperate waters" (Jereb & Roper, 2010) Jereb & Roper, 2010 115 60 "North Atlantic, primarily 20–60°N, including Mediterranean Sea" (Bolstad 2010) (Jereb & Roper, 2010)(Salman et al. 2002) -55 45 N to 55 S (Jereb & Roper, 2010) (Jereb & Roper, 2010) 4 1 “Based on the results of the study it can be assumed that only paralarvae and juvenile stages of A. lichtensteinii, driven by currents and water masses from the Mediterranean, appear in areas of the South Adriatic Pit. This early stages are probably driven from the Levantine basin through Otranto Straits to the Adriatic Sea by currents carrying water masses of the surface and intermediate layer Levantine waters from the Mediterranean (Robinson, Leslie, Theocharis, & Las caratos, 2001; Zore-Armanda, 2000).” (Sifner et al. 2018) (Sifner et al. 2018) 1 "their vertical distribution demonstrate that this species displays vertical migration
at least during their early life stages" (Roura et al., 2019)		 (Roura et al., 2019) southern Adriatic Sea “The life span of A. lichtensteinii was not possible to assess due to the lack of the mature specimens in the sample, but it was found that the oldest individual was approximately 6 months old (178 days) and its mantle length was only 106 mm…When taking into consideration that this species reaches maximum mantle length of 300 mm, that males mature at about 200 mm ML (Kubodera, Piatkowski, Okutani, & Clarke, 1998), and also that with aging the growth rate of individuals slows down, it can be concluded that A. lichtensteinii is a slowgrowing species” (Sifner et al. 2018) (Sifner et al. 2018) southern Adriatic Sea “The life span of A. lichtensteinii was not possible to assess due to the lack of the mature specimens in the sample, but it was found that the oldest individual was approximately 6 months old (178 days) and its mantle length was only 106 mm…When taking into consideration that this species reaches maximum mantle length of 300 mm, that males mature at about 200 mm ML (Kubodera, Piatkowski, Okutani, & Clarke, 1998), and also that with aging the growth rate of individuals slows down, it can be concluded that A. lichtensteinii is a slowgrowing species” (Sifner et al. 2018) (Sifner et al. 2018) NA 0 NA 0 1 Present (Jereb & Roper, 2010) (Jereb & Roper, 2010) 2 "It feeds on epipelagic and upper mesopelagic finfishes and crustaceans" (Jereb & Roper, 2010) Jereb and Roper 2010 0 0 2 7 "is
preyed upon by cetaceans, e.g. Risso’s dolphin and striped dolphins, sperm
whales, and pelagic fishes, e.g. swordfish, and by giant red shrimp in the
Mediterranean" (Jereb & Roper, 2010)		 (Blanco et al., 2006)  (Jereb & Roper, 2010) (Cherel
et al., 2009) (Battaglia
et al. 2013) (Consoli
et al. 2008) (Rasero
et al. 1993)		 3 0 not gregarious (inferred from photo and video material) NA 0 NA 19 12 189.224 154.938 160 98 W-W
Architeuthis dux Architeuthis dux 900 "A specimen of Architeuthis was photographed alive for the first time as it attacked a baited camera off Japan at 900 m." (Jereb & Roper, 2010) (Jereb & Roper, 2010; Kubodera et al., 2018; Wada et al. 2015) 200 "The vertical distribution of Architeuthis remains difficult to assess precisely, but captures in deep-sea fishing nets and more accurate information on the foraging behaviour of its principal predator, sperm whales, indicate a range of about 200 to 800 m (perhaps to 1 000 m). The zone of maximum concentration appears to be about 400 to 600 m." (Jereb & Roper, 2010) (Jereb & Roper, 2010; Kubodera et al., 2018; Wada et al. 2015) 1 "large mesopelagic cephalopods (including Architeuthis)…As with many meso- and bathypelagic squids, giant squid incorporate pockets of ammonia solution within their flesh to enable neutral buoyancy (Clarke et al. 1979)." (Kubodera & Mori 2005). "Slope, seamounts, ?near bottom" (Gomes-Pereira et al., 2016)
"These sources also indicate habitats associated with the bottom as well as in deep midwater layers well above very great bottom depths." (Jereb & Roper, 2010)		 Kubodera & Mori 2005; Gomes-Pereira et al., 2016 120 70 "Normally doesn’t live to the north from the Northern Sea;
single stranding specimens rarely reach norwegian shelf up
to 70°N)" (Xavier et al., 2018) although (Cherel, 2003) writes" This is in agreement with the general view that giant squid are
broadly distributed from 808N in the eastern North
Atlantic to the Subtropical Front at about 408S and in the
Atlantic, Indian and Paci¢c Oceans (Clarke, 1986). The
diet of sharks moreover expands their distribution further south (to about 508S) in colder waters of the Indian Ocean."		 (Xavier et al., 2018) -50 " This is in agreement with the general view that giant squid are
broadly distributed from 808N in the eastern North
Atlantic to the Subtropical Front at about 408S and in the
Atlantic, Indian and Paci¢c Oceans (Clarke, 1986). The
diet of sharks moreover expands their distribution further south (to about 508S) in colder waters of the Indian Ocean." (Cherel, 2003)		 (Cherel, 2003) 4 1 Japan "In particular, the smallest individual (Figure 2) of this study may normally occupy shallow water at night because it was captured alive in the upper depth of 45 m in the early morning where it may have wandered into neritic waters. Guerra et al. (2010) have suggested an ontogenetic shift in diet of the giant squid using stable isotope recorded in beaks. Thus, the giant squids may inhabit and migrate through different depths of the ocean depending on the development stage. Another possibility for the occurrence of the juvenile and young individuals near the coast is that they may have drifted there with warm-water currents. The coasts of Kyushu Island and the south-western Sea of Japan have been known to pass through the Kuroshio and Tsushima Warm Current, respectively (e.g. Akitomo et al., 1997; Hase et al., 1999). It is possible that some individuals in the early life stages of this species had come into the coastal area via offshore flow and the upwelling of warm currents." (Wada et al. 2015) (Wada et al. 2015; Kubodera et al., 2018; Jereb & Roper 2013) 1 "ontogenetic change in diet early in life history, age of at least 2 years, and sedentary behaviour in adulthood, with gradual ontogenetic descent. " (Jereb & Roper, 2010) (Jereb & Roper, 2010; Wada et al. 2015; Winkelmann et al. 2014;) 1 "Japan "" In particular, the smallest individual (Figure 2) of this study may normally
occupy shallow water at night because it was captured alive in
the upper depth of 45 m in the early morning where it may
have wandered into neritic waters. Guerra et al. (2010) have suggested an ontogenetic shift in diet of the giant squid using stable
isotope recorded in beaks. Thus, the giant squids may inhabit
and migrate through different depths of the ocean depending
on the development stage. Another possibility for the occurrence
of the juvenile and young individuals near the coast is that they
may have drifted there with warm-water currents. The coasts of
Kyushu Island and the south-western Sea of Japan have been
known to pass through the Kuroshio and Tsushima Warm
Current, respectively (e.g. Akitomo et al., 1997; Hase et al.,
1999). It is possible that some individuals in the early life
stages of this species had come into the coastal area via offshore
flow and the upwelling of warm currents."" (Wada et al. 2015)"		 (Wada et al. 2015) 5113.5 "As with other cephalopods the growth rate of Architeuthis is thought to be very rapid, and full adult sizes (up to perhaps 500 kg) may be attained in a maximum of 2 to 3 years based on statolyth analysis. However, isotopic analysis using delta super (18) Oxygen suggest a longer life span, perhaps to about 14 years." (Jereb & Roper, 2010) [ 1 – 14 ] (Perales-Raya et al., 2020; Romanov et al., 2018; Kubodera et al., 2018; Jereb & Roper, 2010; ) 365.3 "Geographical variations may influence the life-span duration from different
locations and therefore the maximum ages in the present study from the largest in-
dividual measured (see Table 4) should be taken with caution, as the specimen was
caught off the coast of southern Australia (Norman and Lu 1997). Nonetheless, a
maximum lifespan of around 3 yrs is plausible in the species, in accordance with
previous studies suggesting a longevity of 2 yrs for males and 3 yrs for females based
on modal size frequency distribution of mass findings (Kubodera et al. 2018) and
statolith ages (e.g., Jackson et al. 1991, Fernández-Núñez and Hernández-González
1995, Lordan et al. 1998) in both the Pacific Ocean and the Atlantic Ocean. Guerra
et al. (2006) summarized the age estimation of A. dux from different approaches and
suggested lifespan durations between 1 and 3 yrs. They also found a marked sexual
dimorphism in A. dux and suggested that males have a lifecycle of around 1 yr and
are much smaller than females. Females are larger, mature later, and live for 2 to 3
yrs. Regarding those authors, it seems unlikely that females reach greater ages. The
results of the present study are not in line with long life-span estimations of many
years from isotopic analysis on statoliths of three specimens from Tasmanian waters
(Landman et al. 2004). Ages derived from the model fell between 13 and 38 yrs, with
<14 yrs cited as being most likely; however, the parameters are extremely dependent
on assumptions of depth and statolith growth patterns, making the age estimates
highly uncertain (Roper and Shea 2013)." (Perales-Raya et al., 2020)		 (Perales-Raya et al., 2020; Romanov et al., 2018; Kubodera et al., 2018; Jereb & Roper, 2010; ) 422 "If correct this would make them
one of the fastest growing animals in
the world. Three mature males caught
in Ireland were about half the size of
this specimen and studies of their
statoliths suggested ages of 294 to
422 days (10 to 14 months). It may be
that males mature smaller and
younger than females. " (Herdson, 2002)		 (Herdson, 2002) 294 "If correct this would make them
one of the fastest growing animals in
the world. Three mature males caught
in Ireland were about half the size of
this specimen and studies of their
statoliths suggested ages of 294 to
422 days (10 to 14 months). It may be
that males mature smaller and
younger than females. " (Herdson, 2002)		 (Herdson, 2002) 3 3 1 "The statocysts of Architeuthis are oblique to the body axis (Roeleveld and Lipiński 1991), suggesting that the animal orients obliquely in the water column. This observation supports the hypothesis that giant squid hang vertically or in a slightly head-down position in the water column to capture prey species schooling below, such as the epibenthic orange roughy around New Zealand seamounts. The observation of red pigmentation of the inner mantle surface (Fig. 6A) suggests the hypothesis that Architeuthis eats biolumiscent prey" (Roper & Shea 2013) Roper & Shea 2013
Regueira et al. 2014		 1 "Midwater fishes with tissues that are high in protein and low in water content often are strong swimmers with high metabolism (Childress et al. 1980), and Architeuthis fi ts this pattern (Robison 1989). Recent video of a jig-captured specimen strongly ejecting water through its funnel (Kubodera 2010) suggested that the giant squid is a strong swimmer. The functionality of the tentacles in prey capture was similarly debated until Kubodera and Mori (2005) showed that the tentacles are used to strike at prey in a manner consistent with other active hunters (Messenger 1968, Kier 1982)." (Roper & Shea 2013) Roper & Shea 2013 1 "Although prey items must be well masticated by cephalopods in order to pass through the narrow oesophagus (maximum relaxed diameter 10 mm in this specimen), one prey fragment recovered from the stomach caecum was of 69 mm greatest dimension, and 19 mm compressed dimension; many other fragments were of comparable size. It appears that Architeuthis dispatches prey by slicing it into large pieces and passing them down the oesophagus. Nevertheless, the dimensions of larger prey items are at striking odds to the diameter of the relaxed oesophagus, especially given that the oesophagus in cephalopods passes directly through the brain." (Bolstad & O´Shea, 2004) Bolstad & O'Seah 2004 NA 0 1 (Jereb & Roper, 2010) 21 "Prey of Architeuthis consist of macrourid fishes (including Macruronus novaezelandiae, the blue grenadier), the blue whiting (Micromesistius poutassou), squids (including Architeuthis, Nototodarus) and orange roughy (Hoplostethus atlanticus). Analysis of tracers, heavy metals and stable isotopes suggests that Architeuthis is a very high level predator, feeding on high trophic level fishes and squids. Stable isotope and trace element analyses of Architeuthis beaks indicate aspects of its biology: ontogenetic change in diet early in life history, age of at least 2 years, and sedentary behaviour in adulthood, with gradual ontogenetic descent." (Jereb & Roper, 2010) Jereb & Roper, 2010
Xavier et al., 2018
Bolstad & O´Shea, 2004
Guerra and Rocha 1994
Katsanevakis et al 2008		 0 1 Jereb & Roper, 2010
Bolstad & O´Shea, 2004
Elsevier, 2014		 5 7 "Sperm whale; greenland shark; large pelagic osteichthyes" (Xavier et al., 2018) Voss, 1956
Jereb & Roper, 2010
Xavier et al., 2018
Romanov et al., 2018
Cherel, 2003
Laptikhovsky et al. 2020		 4 1 "Stranding records and deep trawling captures suggest Architeuthis is a solitary animal." (Roper & Shea 2013); "Aldrich (1991, p. 475) attributes this damage to combat with another architeuthid. Our report of large tentacular sucker rings of Architeuthis from a stomach caecum, and that of Zeidler & Gowlett-Holmes (1996), of large unidentifiable suckers recovered from a conspecific stomach, equally support Aldrich's contention that damage through combat with other architeuthids is possible." [Found in a mature female squid] (Bolstad & O´Shea, 2004) (Roper & Shea 2013) (Bolstad & O´Shea, 2004) 0 "Stranding records and deep trawling captures suggest Architeuthis is a solitary animal." (Roper & Shea 2013); "Aldrich (1991, p. 475) attributes this damage to combat with another architeuthid. Our report of large tentacular sucker rings of Architeuthis from a stomach caecum, and that of Zeidler & Gowlett-Holmes (1996), of large unidentifiable suckers recovered from a conspecific stomach, equally support Aldrich's contention that damage through combat with other architeuthids is possible." [Found in a mature female squid] (Bolstad & O´Shea, 2004) (Roper & Shea 2013) (Bolstad & O´Shea, 2004) 1 0 1 1 “carcass of a giant squid had stranded on Bares beach…it showed numerous sucker marks (diameter 2.14–4.60 cm) and scrapes on the skin (Fig. 1); (6) the teeth of the sucker rings were short and of a similar size; and (7) the specimen had a long (24 cm)…Based on a beached specimen with sucker scars on its mantle and arms (2.7–4.0 cm diameter), and serrations on some arms inconsistent with damage caused by beaching, Aldrich (1991) attributed these injuries to combat with a conspecific. Sucker scars on the Bares specimen have comparable diameter to those reported by Aldrich (1991), suggesting a similar battle, apparently with another large cephalopod. The beak of the other animal might have produced the deep scratches in the skin…The victim was an immature unmated female, which seems to exclude competition for mates. Cannibalism is also unlikely due to the absence of large bite marks. The only other cephalopod species in the area of sufficient size to fight a squid as big as the Bares specimen are Ommastrephes bartramii or Taningia danae. However, the shape of the sucker rings of O. bartramii is not consistent with the marks observed in the skin of the Bares specimen and in T. danae the suckers are transformed into hooks. Therefore, we suggest that the Bares specimen was engaged in battle with a conspecific.” (Guerra et al. 2018) & "Aldrich (1991, p. 475) attributes this damage to combat with another architeuthid. Our report of large tentacular sucker rings of Architeuthis from a stomach caecum, and that of Zeidler & Gowlett-Holmes (1996), of large unidentifiable suckers recovered from a conspecific stomach, equally support Aldrich's contention that damage through combat with other architeuthids is possible." [Found in a mature female squid] (Bolstad & O´Shea, 2004) (Guerra et al. 2018) 51 33 1766.6 1570 NY
Argonauta argo Argonauta argo 1500 " found in the surface
layer both day and night but also down to 1500 m deep." (Judkins & Vecchione, 2020)		 (Lin-lin et al., 2015; Finn & Norman, 2010; Tejerina, 2019; Maximiliano et al., 2015; Cairns, 1976; Stevens et al., 2015; Judkins & Vecchione, 2020) 0 "Finn and Norman (2010) observed that female argonauts
trap air inside the shell at the sea surface to acquire neu-
tral buoyancy. This method is thought to be effective only
at very shallow water depths of <10 m (Finn and Norman
2010). Female argonauts are therefore restricted to the
uppermost 10 m of the surface waters" (Stevens et al., 2015)		 (Lin-lin et al., 2015; Finn & Norman, 2010; Tejerina, 2019; Maximiliano et al., 2015; Cairns, 1976; Stevens et al., 2015; Judkins & Vecchione, 2020) 1 "Males might live inside pelagic salps." (Gomes-Pereira et al., 2016) Gomes-Pereira et al., 2016 90 45 The northernmost record of stranded paper nautiluses in Japan is at 45 degrees north (Funaki & Sato, 2009) (Funaki & Sato, 2009) -45 "cosmopolitan distribution in tropical and subtropical seas between approx. 45°N and 45°S (Roper et al. 1984; Finn 2013)" (Stevens et al., 2015) (Stevens et al., 2015) 1 0 35 individuals caught "There is no evidence of ontogenic shift." (Judkins & Vecchione, 2020) (Judkins & Vecchione, 2020) 0 35 individuals caught "There is no evidence of ontogenic shift." (Judkins & Vecchione, 2020) (Judkins & Vecchione, 2020) "So far, their exact growth rate and lifespan are unknown." (Stevens et al., 2015) "It is quite doubtful that Argonauta lives more than a few years." (Nishimura, 1968) 3 3 1 "The pelagic octopods such as Argornauta and small Tremocropus are also touch feeders [i.e., tactile feeding]. In both genera, there is a well-developed web (Fig. 4.7). Young (l960a) describes how in Argonauta this extends from the first arm and can be spread over the thin shell; when food touches the web, the fourth arm sweeps out to seize it and pass it to the mouth. Young never saw the animal attack prawns or fish in the aquarium. These animals are slow-moving creatures with relatively weak suckers, and they presumably feed on the plankton among which they drift; in the sea they are known to feed on pteropods (Okutani, 1960]." (Hanlon & Messenger, 2018, p. 82) Hanlon and Messenger 2018 1 "Active attacks to seize prey, as reported for cuttlefish and squids, have never been observed for argonauts." (Heeger et al., 1992) Heeger et al., 1992
Bello & Rizzi, 1990
Young, 1960		 1 "A female found holding a jellyfish tightly, with the suckers of the lateral and ventral arms, was watched by scuba divers for an hour (Heeger er al. 1992). Both animals were collected and when placed in an aquarium the argonaut ejected itself from the medusa. Large pieces of mesoglea were missing, and a few distinct holes in the centre of the bell of the medusa were interpreted as bites. Argonauts apparently feed, take shelter, and even obtain a free passage from other animals." (Nixon & Young 2003) Nixon and Young 2003
Heeger et al., 1992
Nixon & Young 2003		 5 5 1 "Banas et al. (1982) observed juvenile argonauts inside the branchial cavity of salp chains. Although many zooplankton organisms were in association with the salp chains, the argonauts had neither fed on them nor had they damaged the salp tissue which the authors documented by examination of gut contents. Banas et al. concluded that this association provided flotation, transportation or camouflage to the argonauts. " (Heeger et al., 1992) Heeger et al., 1992 1 "When alarmed , the argonaut is capable of putting on an impressive color display. Within two-thirds of a second, it can turn dark red" (Iliffe, 1982 Iliffe, 1982
Young, 1960		 1 In addition, its color pattern, called countershading (dark shades on the upper surface and reflective silver on the underside), serves as excellent camou Rage for an animal swimming near the surface of the sea." (Iliffe, 1982) Iliffe, 1982 1 "In Octopus vulgaris, the Deimatic Display comprises six components: (1) paling of the skin; (2) arms curved in wide arc and web spread maximally; (3) Dark eye ring, (4) Dilated pupil; (5) Dark edged suckers to create a dark margin to the octopus’s outline; and (6) jetting water. Deimatic Displays in other octopuses differ slightly. For example, Octopus maya has diffuse ‘spots’ but no distinct ocelli and Octopus bimaculoides, O. bimaculatus and O. filosus have prominent ocelli at the bases of their arms that are used with lightly graded mottles to produce a Deimatic Display. Quite different Deimatic Displays are shown by Octopus macropus and Octopus ornatus. The arms or web are spread to enlarge the appearance of the body, and there is maximal contrast of light/dark on the body: both species turn bright reddish-brown with bright white ovals all over the body. The deimatic response in the oceanic octopod Argonauta argo (the paper nautilus) is different again and occurs when the brightly silver iridescent web (part of the first arm pair) that covers the shell is quickly withdrawn by the octopus (Young, Reference Young1960a)." (Hanlon & Messenger, 2018, p. 125-126) Hanlon & Messenger, 2018, p. 125-126 1 Inking (Heeger et al., 1992) Heeger et al., 1992
Iliffe, 1982
Young, 1960		 1 Present in the family (Jereb et al., 2014) (Jereb et al., 2014) 4 "According to Nesis (1977), the natural diet of female argonauts consists mainly of heteropods and pteropods. Robson (1932) described Argonauta spp. preying upon small fish and crustaceans. The symbiosis of juvenile argonauts with salps may also provide a food source for the cephalopod (Banas et al. 1982). " (Heeger et al., 1992) Heeger et al 1992
Nabhitabhata et al 2009
Corsini-foka et al., 2011		 0 0 3 9 "argonauts are frequently found as stomach contents of epipelagic predators, e.g., billfishes, tunas and sharks (e.g., Roper et al. 1984; Staudinger et al. 2013)." (Stevens et al., 2015) Randall et al., 1981
Ribeiro Simoes & Andrade, 2000
Ortiz-Corps et al., 1995
Manooch et al. (1984)
Tejerina, 2019
Stevens et al., 2015
Corsini-foka et al., 2011
Blanco et al., 2006
Kousteni et al. 2018
Bello 1997		 5 2 "The two argonauts kept in the same tank did not show any sign of suffering, contrary to what happens to many benthic octopods, which, when in a confined space, may even display cannibalism." (Bello & Rizzi, 1990) (Bello & Rizzi, 1990) 0 "The two argonauts kept in the same tank did not show any sign of suffering, contrary to what happens to many benthic octopods, which, when in a confined space, may even display cannibalism." (Bello & Rizzi, 1990) (Bello & Rizzi, 1990) 2 0 2 1 "Similar to observations of A. nouryi by Rosa and Seibel (2010), A. hians in the Andaman Sea (Sukhsangchan et al., 2009), and A. argo in the Mediterranean Sea (Salman, 2004), our results suggest aggregative spawning behaviors also occur in the Northwest Atlantic." (Staudinger et al. 2012) (Staudinger et al. 2012) 1 "females brood the eggs in a specific ‘‘shell’’, or pseudo-conch until hatching" (Laptikhovsky & Salman, 2002) Laptikhovsky & Salman, 2002 0 "females brood the eggs in a specific ‘‘shell’’, or pseudo-conch until hatching" (Laptikhovsky & Salman, 2002) Laptikhovsky & Salman, 2002 103 45 121.66 86.548 34.6 50 40 W-W-NY
Bathothauma lyromma Bathothauma lyromma 2000 "Its vertical range extends from subsurface epipelagic depths as paralarvae and undergoes very clear ontogenetic descent through the
mesopelagic zone and into the bathypelagic waters to in excess of 2 000 m. " (Jereb & Roper, 2010)		 (Jereb & Roper, 2010; Dilly and Herring 1974) 200 "Allan (1940) reported a juvenile B. lyromma taken at the surface and Voss (1960) recorded a juvenile taken at 200 m. All other records are from open nets fished in excess of 520 m. " (Cairns, 1976; Dilly and Herring 1974; Aldred 1974) (Jereb & Roper, 2010; Dilly and Herring 1974) 1 "This species was found throughout the water column during the day and deeper depths (below 600 m) at night. A weak ontogenic shift was noted for this species but more material is required to confirm this pattern" (Judkins & Vecchione, 2020) Cairns, 1976
Judkins & Vecchione, 2020		 85 45 45 N to 40 S (Jereb & Roper, 2010) (Jereb & Roper, 2010) -40 45 N to 40 S (Jereb & Roper, 2010) (Jereb & Roper, 2010) 3 1 "Its vertical range extends from subsurface epipelagic depths as paralarvae and undergoes very clear ontogenetic descent through the
mesopelagic zone and into the bathypelagic waters to in excess of 2 000 m. " (Jereb & Roper, 2010)		 (Jereb & Roper, 2010; Judkins & Vecchione, 2020) NA 0 NA 0 0 0 2 Found in the stomach content of Cook's Petrel Pterodroma cookii (Imber, 1996) Imber 1978
Imber 1996		 1 0 not gregarious (inferred from photo and video material) NA 0 NA 7 5 46.4 165 NY
Bathypolypus bairdii Bathypolypus bairdii 1545 "Depths range from 20 to 1 545 m. " (Jereb et al., 2014) (Roura et al 2010; Vecchione & Roper, 1991, p. 438; Jereb et al., 2014) 20 "Depths range from 20 to 1 545 m. " (Jereb et al., 2014) (Roura et al 2010; Vecchione & Roper, 1991, p. 438; Jereb et al., 2014) 2 inferred from ROV data (see e.g., https://oceanexplorer.noaa.gov/okeanos/explorations/ex1806/logs/june29-2/june29-2.html) inferred from ROV data (see e.g., https://oceanexplorer.noaa.gov/okeanos/explorations/ex1806/logs/june29-2/june29-2.html) 5 50 "Distribution Saguenay Fjord,
southern Gaspe waters (Baie des Chaleurs, Gaspe Bay to American, Orphan
and Bradelle banks; eastern boundary: eastern Bradelle Valley),
downstream part of middle St. Lawrence estuary, lower St. Lawrence
estuary, Laurentian Channel (bathyal zone)(=Honguedo Strait); south
Slope of Anticosti Island; Cobscook Bay" Taken from WoRMS 2023-04-05		 Taken from WoRMS 2023-04-05 45 "Distribution Saguenay Fjord,
southern Gaspe waters (Baie des Chaleurs, Gaspe Bay to American, Orphan
and Bradelle banks; eastern boundary: eastern Bradelle Valley),
downstream part of middle St. Lawrence estuary, lower St. Lawrence
estuary, Laurentian Channel (bathyal zone)(=Honguedo Strait); south
Slope of Anticosti Island; Cobscook Bay" Taken from WoRMS 2023-04-05		 Taken from WoRMS 2023-04-05 4 1095.8 "Mortality must be very low. O'Dor & Macalaster (1983) show that a three-year lifespan,
including one reproduction period, is probable, but that a longer life cannot be ruled out." (Muus 2002)		 (Muus 2002) 1095.8 "Mortality must be very low. O'Dor & Macalaster (1983) show that a three-year lifespan,
including one reproduction period, is probable, but that a longer life cannot be ruled out." (Muus 2002)		 (Muus 2002) 2 2 1 "Based on aquaria observations, O'Dor & Macalaster (1983) suggest that bairdii practises a sit-and-wait feeding strategy, and they list food items demonstrating its omnivorous nature. Undenwater photos confirm the supposed feeding strategy. In the prawn trawling grounds, bairdii is surrounded by a rich food supply of roaming crustaceans, polychetes, and molluscs." (Muus 2002) Muus 2002 1 "Voight (2008) suggested that Benthoctopus and Graneledone may forage on infauNAby sweeping the mid section of their arms through the sediment [i.e., tactile feeding]." (Hanlon & Messenger, 2018, p. 81) Hanlon & Messenger, 2018 NA 0 0 no ink sac in the genus (Jereb et al., 2014)
(Bello, 2004; Jereb et al., 2014)		 0 Absent in the genus (Jereb et al., 2014) (Jereb et al., 2014) 0 0 0 not gregarious (inferred from photo and video material) 1 0 1 0 "To deposit and guard the eggs, the females need a firm substratum. […] Hiding egg-guarding females are less apt to be caught in a trawl. " (Muus 2002) Muus 2002 1 "To deposit and guard the eggs, the females need a firm substratum. […] Hiding egg-guarding females are less apt to be caught in a trawl. " (Muus 2002) Muus 2002 6 6 193 89 NY
Bathypolypus sponsalis Bathypolypus sponsalis 1835 "This species inhabits depths down to 1835 m in the Mediterranean Sea, with main distribution between 400 and 700 m (Villanueva, 1992)" (Salman et al., 2001) (Salman et al., 2001; Quetglas et al., 2000; Jereb et al., 2014; Cuccu et al 2011; Voliani et al 2009; Roura et al 2010; Quetglas et al 2001; D'Onghia et al 1993; Villanueva 1992; Muus 2002) 170 "170-1250 m" (Muus 2002) (Salman et al., 2001; Quetglas et al., 2000; Jereb et al., 2014; Cuccu et al 2011; Voliani et al 2009; Roura et al 2010; Quetglas et al 2001; D'Onghia et al 1993; Villanueva 1992; Muus 2002) 2 "Bathypolypus sponsalis was found in both areas on muddy bottoms" (D'Onghia et al 1993) D'Onghia et al 1993 30 45 "Bathypolypus sponsalis (Fischer and Fischer, 1892) has been reported from the Bay of Biscay to Cape Verde in the Atlantic Ocean and also in the MediterraeNASea (Voss 1988a; Guerra 1992)." (Quetglas et al 2001). (Quetglas et al 2001). 15 "Bathypolypus sponsalis (Fischer and Fischer, 1892) has been reported from the Bay of Biscay to Cape Verde in the Atlantic Ocean and also in the MediterraeNASea (Voss 1988a; Guerra 1992)." (Quetglas et al 2001). (Quetglas et al 2001). 4 1 "Small individuals occur at greater depths than larger individuals, suggesting up-slope ontogenetic migration." (Jereb et al., 2014) (Jereb et al., 2014; Villanueva 1992) 1461 Life cycle: not known, but closely related species B. arcticus completes its life cycle in 2-3 years (excluding embryo stage; Nixon & Young, 2003); "assuming daily deposition of stylet increments, specimens of the deepwater octopod Bathypolypus sponsalis were estimated to be <1 year old (Barratt and Allcock, 2010), which does not agree with the longer lifespan (3–4 years) suggested by laboratory-kept individuals (O’Dor and Malacaster, 1983)" (Hoving et al, 2014:308, in Vidal, ed.) (Hoving et al, 2014:308, in Vidal, ed.) 365.3 Life cycle: not known, but closely related species B. arcticus completes its life cycle in 2-3 years (excluding embryo stage; Nixon & Young, 2003); "assuming daily deposition of stylet increments, specimens of the deepwater octopod Bathypolypus sponsalis were estimated to be <1 year old (Barratt and Allcock, 2010), which does not agree with the longer lifespan (3–4 years) suggested by laboratory-kept individuals (O’Dor and Malacaster, 1983)" (Hoving et al, 2014:308, in Vidal, ed.) (Hoving et al, 2014:308, in Vidal, ed.) 517.4 Males 4-6 months and females 8-17 months with Mangold-Wirz
(1963) as reference in (Nixon, 1969)		 (Nixon, 1969) 121.8 Males 4-6 months and females 8-17 months with Mangold-Wirz
(1963) as reference in (Nixon, 1969)		 (Nixon, 1969) NA 0 NA 0 0 no ink sac in the genus (Jereb et al., 2014)
(Bello, 2004; Jereb et al., 2014)		 0 Absent in the genus (Jereb et al., 2014) (Bello, 2004; Jereb et al., 2014) 19 "Stomach content analysis yielded a total of 19 different prey items belonging to four major groups (Crustacea, Mollusca, Ophiuroidea and Osteichthya). (Quetglas et al 2001). Quetglas et al 2001
Nixon & Young 2003		 0 1 Quetglas et al 2001 4 3 Found in the stomach of the Mediterranean monk seal, Monachus monachus (Salman et al., 2001) Salman et al., 2001
Villanueva 1992		 2 0 not gregarious (inferred from photo and video material) NA 0 NA 20 10 78.376 51.992 38 45 30 25 W-W-NY
Bathyteuthis abyssicola Bathyteuthis abyssicola 4200 "An oceanic species that occurs between
about 100 and 4 200 m depth, but normally it is encountered
between 700 and 2 000 m in the Southern Ocean where it carries
out a deep vertical diurnal migration. Paralarvae and juveniles
tend to live at shallower depths than the adults." (Jereb & Roper, 2010)		 (Jereb & Roper, 2010; Young, 1972; Roper 1969; Boyle & Rodhouse, 2005) 500 [AL: Due to juveniles and paralarvea tends to live at shallower depth, data for adults are uncertain. Based on the different observations it seems fair to put it at 500 min depth] "" abyssicola has been intensively studied in Antarctic waters where it seldom has been captured in depths less than 500 m; it shows a peak occurrence at depths of 1000 to 2500 m (ROPER, 1969). Off Hawaii a single capture has been made between 830 and 975 m in an opening–closing trawl." (Young, 1972)" But see "An oceanic species that occurs between
about 100 and 4 200 m depth, but normally it is encountered
between 700 and 2 000 m in the Southern Ocean where it carries
out a deep vertical diurnal migration. Paralarvae and juveniles
tend to live at shallower depths than the adults." (Jereb & Roper, 2010)		 (Jereb & Roper, 2010; Young, 1972; Roper 1969; Boyle & Rodhouse, 2005) 1 "Oceanic, mesop. " (Gomes-Pereira et al., 2016) Gomes-Pereira et al., 2016 130 50 "Bathyteuthis abyssicola, Hoyle, a young specimen of which
occurred at 50 ° 22' N., 11 ° 40' W." (Massy 1916)		 (Massy 1916) -80 45 N to 80 S (Jereb & Roper, 2010) (Jereb & Roper, 2010) 4 1 "An oceanic species that occurs between
about 100 and 4 200 m depth, but normally it is encountered
between 700 and 2 000 m in the Southern Ocean where it carries
out a deep vertical diurnal migration. Paralarvae and juveniles
tend to live at shallower depths than the adults." (Jereb & Roper, 2010)		 (Jereb & Roper, 2010) 1 "An oceanic species that occurs between
about 100 and 4 200 m depth, but normally it is encountered
between 700 and 2 000 m in the Southern Ocean where it carries
out a deep vertical diurnal migration. Paralarvae and juveniles
tend to live at shallower depths than the adults." (Jereb & Roper, 2010)		 (Jereb & Roper, 2010) NA 0 NA 0 "NA" (Xavier et al., 2018) Xavier et al., 2018) 0 0 2 "Among predators of B. abyssicola are the melon-headed whale in Hawaiian waters." (Jereb & Roper, 2010) Jereb & Roper, 2010 1 0 not gregarious (inferred from photo and video material) 1 0 1 “southern Kerguelen Plateau…Squids at stages 0 to I were predominant (ML < 100 mm), with a single size mode for each species, suggesting that these species may use the plateau as a spawning and/or nursery ground.” (Lin et al. 2020) 16 13 7 30 NY
Bolitaena pygmaea Bolitaena pygmaea 1600 Caught between 1050 – 1600 m by Urbano & Hendrickx (2018) (Judkins & Vecchione, 2020; Jereb et al., 2014; Urbano & Hendrickx, 2018) 0 "Seventy-seven BolitaeNApygmaea (6–64 mm ML) are distributed
from 0 to 1500 m with no diel migration pattern observed." (Judkins & Vecchione, 2020)		 (Judkins & Vecchione, 2020; Jereb et al., 2014; Urbano & Hendrickx, 2018) 1 "These small pelagic octopuses typically live over deeper water." (Jereb et al., 2014) Jereb et al., 2014 88 45 45 N to 43 S (Jereb et al., 2014) (Jereb et al., 2014) -43 45 N to 43 S (Jereb et al., 2014) (Jereb et al., 2014) 3 1 "This species is an upper-boreal, meso-bathypelagic species that undergoes ontogenetic descent and diel vertical migration." (Jereb & Roper, 2010) (Jereb & Roper, 2010; Judkins & Vecchione, 2020; Jereb et al., 2014) 1 "This species is an upper-boreal, meso-bathypelagic species that undergoes ontogenetic descent and diel vertical migration." (Jereb & Roper, 2010) (Jereb & Roper, 2010) NA 0 NA 0 0 0 3 "Northern fur seal; alaska pollock" (Xavier et al., 2018) Tsuchiya and Sawadaishi 1997
Xavier et al., 2018		 2 0 not gregarious (inferred from photo and video material) 2 0 2 "Pigmentation greatly increases in females as they mature and the arms become relatively longer. Increased pigmentation may be associated with the need to mask output from the female’s circumoral light organ. This light organ may be used for reproductive signalling to males. " (Jereb et al., 2014) 1 "Females brood the eggs by holding the egg mass within their arm crown" (Jereb et al., 2014) Jereb et al., 2014 0 "Females brood the eggs by holding the egg mass within their arm crown" (Jereb et al., 2014) Jereb et al., 2014 2 2 28.8 75 NY
Callistoctopus macropus Callistoctopus macropus 242 Depth range (m): 102–242 (Sifner et al., 2005) (Sifner et al., 2005; Voss & Phillips 1957) 10 Azores "Both specimens examined here were caught in the littoral by divers (<10 m) (Goncalves 1991) (Sifner et al., 2005; Voss & Phillips 1957) 2 "occur on a range of habitats from soft sediments (sand and mud) and seagrass beds, to coral and rocky reefs" (Norman & Sweeney 1997) Lykkeboe & Johansen, 1982
Norman & Sweeney 1997		 32 45 45 N to 13 N (Jereb et al., 2014) (Jereb et al., 2014) 13 45 N to 13 N (Jereb et al., 2014) (Jereb et al., 2014) 1 2 2 1 " Octopus maci-opus also inserts its very long arms into holes in the sand to extract crustaceans (Hochberg & Couch, 1971; Hanlon er al., 1979)." (Hanlon & Messenger, 2018, p. 81) Hanlon & Messenger, 2018 1 Drilling is well documented (Hiemstra, 2015) Hiemstra, 2015
Fujita, 1916
Ambrose et al., 1988		 5 3 1 captured female in lab "Of the available hiding places, the animal chose a horizontal, externally blackened perspex cylinder 19 cm in diameter, 40 cm in legnth and closed on one end" (Boletzky et al. 2001) Boletzky et al. 2001 1 O. macropus and Macrotritopus defilippi make narrow deep holes in sand (Hanlon, 1988; Hanlon, Hanlon, Watson and Barbosa 2010) Hochberg and Couch1971
Hanlon1988;
Hanlon, Watson and Barbosa2010)		 1 captive individual “It is interesting to note that the octopus paid little attention to the presence of the small moray eels, a normally important predator upon octopods. Should one of them approach too closely, it merely fended it off with one of its arms. It is possible that only large moray eels are predators on octopods and that no reaction was forthcoming from small ones” (Voss & Phillips 1957) Voss & Phillips 1957
Hanlon & Messenger, 2018, p. 125		 1 (Jereb et al., 2014) 1 captured individuals in lab fed on crabs (Carcinus maenas) (Boletzky et al. 2001) Boletzky et al 2001 0 0 1 1 "Risso’s dolphin, Grampus griseus (Cuvier, 1812)" (Blanco et al., 2006) Blanco et al., 2006 1 0 not gregarious (inferred from photo and video material) 1 0 1 captive individual “Other specimens in the tank included from time to time small specimens of O. briareus and about 12 small moray eels of the genera Gymno thorax and Echidna…The presence of the small octopuses caused a slightly greater negative response or retreat, but they in turn were highly disinclined to invade the territory of the octopus. No fighting or other direct interplay of action between these two species was noted, as in general they kept strictly out of each other's way” (Voss & Phillips 1957) (Voss & Phillips 1957) 0 lab, female laid eggs and "brooded them till hatching" (Boletzky et al. 2001) (Boletzky et al. 2001) 1 lab, female laid eggs and "brooded them till hatching" (Boletzky et al. 2001) (Boletzky et al. 2001) 63 21 116.4 NY
Chiroteuthis veranii Chiroteuthis veranii 978 southern Levant, E Mediterranean 978m (Paz et al. 2018) (Quetglas et al., 2000; Quetglas et al. 2013; Paz et al. 2018; Vecchione & Roper, 1991, p. 437) 332 W Mediterranean "Two individuals were taken at 332-340 m depth and the remaining five between 609 and 855 m" (Quetglas et al. 2013) (Quetglas et al., 2000; Quetglas et al. 2013; Paz et al. 2018; Vecchione & Roper, 1991, p. 437) 1 "A mesopelagic to bathypelagic species as adults," (Jereb & Roper, 2010) Jereb & Roper, 2010 130 65 65 N to 65 S (Jereb & Roper, 2010) (Jereb & Roper, 2010) -65 65 N to 65 S (Jereb & Roper, 2010) (Jereb & Roper, 2010) 4 1 "A mesopelagic to bathypelagic
species as adults," & "paralarvae and juveniles are epipelagic to mesopelagic " (Jereb & Roper, 2010)		 (Jereb & Roper, 2010) 1 1 1 "It would be interesting to theorize on the value of the peculiarly arranged tentacular club and the large terminal light organ. In an animal apparently poorly suited for swimming, and living in the darkness of the deep sea, it would appear that the extremely long tentacular stalks might be for lowering the long, well armed clubs below the body where the luminous organs would attract planktonic animals within reach of the tangle of hooked suckers which in turn might act like a "jig" used by fishermen. Such an adaptation might easily be a very efficient food catcher and no other explanation seems plausible." (Voss, 1956) Voss, 1956 1 1 1 Chiroteuthis paralarvae "Observations show that the fins undulate slowly, and to escape the squid turns downwards to jet rapidly away, leaving behind it a long, thin, pseudomorph of ink, approximately its own size (Vecchione er al. 1992).” (Nixon & Young 2003) Nixon & Young 2003 1 inc sac with two photophores (Guerra et al. 2007) (Guerra et al. 2007) 3 "Pelagic small crustaceans, mollucs, fish, Kubota et al. (1981) Kubota et al 1981 0 0 3 11 Found in the stomach of the northern bottlenose whale (Hyperoodon ampullatus) (Fernandez et al., 2014) (Fernandez et al., 2014) (Jereb & Roper, 2010) (Xavier et al., 2018) (Beasley et al. 2013) (Cherel & Duhamel, 2004) (Salman & Karakulak 2009) (Öztürk et al. 2007) (Laptikhovsky et al. 2020) (Velez-Rubio et al. 2015) (Kovacˇić et al. 2010) (Cherel et al. 2004) 5 3 "It has been hypothesized that the species is gregarious, because frequently in deep net tows numerous specimens are captured together, generally an uncommon characteristic with deep sea squids." (Jereb & Roper, 2010) (Jereb & Roper, 2010) 1 "It has been hypothesized that the species is gregarious, because frequently in deep net tows numerous specimens are captured together, generally an uncommon characteristic with deep sea squids." (Jereb & Roper, 2010) (Jereb & Roper, 2010) NA 0 NA 29 19 50.2 68 NY
Chtenopteryx sicula Chtenopteryx sicula 1200 "They occupy depths from the surface to 1200 m with many inhabiting the upper mesopelagic zone." (Judkins & Vecchione, 2020) (Quetglas et al., 2000; Judkins & Vecchione, 2020; Quetglas et al. 2013; Roper 1974; Villanueva 1992; Krstulovic Sifner et al 2014; Vidal et al 2010; Vafidis et al 2008) 150 [AL: Paralarvae higher up but no evidence of adults higher than 150] "in Mediterranean "Ctenopteryx sicula is very restricted in vertical range, limited to the upper 150 m" (Roper 1974)" "Paralarvae epipelagic; adults descend to mesopelagic and bathypelagic depths, undergo strong diel vertical migrations" (Jereb & Roper, 2010) (Quetglas et al., 2000; Judkins & Vecchione, 2020; Quetglas et al. 2013; Roper 1974; Villanueva 1992; Krstulovic Sifner et al 2014; Vidal et al 2010; Vafidis et al 2008) 1 "Oceanic, mesopelagic " (Gomes-Pereira et al., 2016) Gomes-Pereira et al., 2016; Jereb and Roper, 2010; Krstulovic Sifner et al., 2014; 95 55 "The specimens were collected on the Amsterdam Mid North Atlantic Plankton Expeditions (1980–1983) along a transect from 55N to 25N along 30W." (Shea and Vecchione 2002) (Shea and Vecchione 2002) -40 45 N to 40 S (Jereb & Roper, 2010) (Jereb & Roper, 2010) 2 1 "the center of abundance of C. sicula appears to shift from the Tyrrhenian Sea in the summer eastward to the Ionian Sea in winter.” (Roper 1974) (Roper 1974) 1 "Paralarvae epipelagic; adults descend to mesopelagic and bathypelagic depths, undergo strong diel vertical migrations" (Jereb & Roper, 2010) (Jereb & Roper, 2010; Judkins & Vecchione, 2020) 1 "Paralarvae epipelagic; adults descend to mesopelagic and bathypelagic depths, undergo strong diel vertical migrations" (Jereb & Roper, 2010) (Jereb & Roper, 2010) NA 0 NA 0 1 "They have light organs (photophores) on the
ink sac" (Norman and Reid 2000 – Book, p47)		 (Norman and Reid, 2000; (Krstuolvic Sifner et al., 2014) 4 "waters off the Balearic Islands..in July. A single stomach from the largest female was analysed (FWI=0.08%), which contained three individuals of the natantian crustacean Gennades elegans." (Quetglas et al. 2013) Vafidis et al 2009 0 0 Quetglas et al. 2013
Vafidis et al 2009		 1 1 Risso's dolphin off Turkish coast (Öztürk et al. 2007) Öztürk et al. 2007 1 0 not gregarious (inferred from photo and video material) NA 0 NA 12 12 24.2 36 NY
Cirroteuthis muelleri Cirroteuthis muelleri 4854 "Depth range from 500 to 4 854 m, with highest
abundance around 3 000 to 3 500 m." (Jereb et al., 2014)		 (Jereb et al., 2014) 500 "Depth range from 500 to 4 854 m, with highest
abundance around 3 000 to 3 500 m." (Jereb et al., 2014)		 (Jereb et al., 2014) 2 "This […] suggests that Cirroteuthidae spend a significant portion of their time drifting in the pelagic realm. While we cannot completely exclude that C. muelleri opportunistically consumes food in the water column, the body morphology of Cirroteuthidae seems not capable of engulfment of prey in the water column, since their suckers are weak and modified […]. This suggests C. muelleri feeds primarily on the seafloor, and will have to go all the way down in order to hunt. This extensive pelagic–benthic migration of thousands of metres to the seafloor is unique among cephalopods." Golikov et al. (2023) Golikov et al. 2023; Jereb et al., 2014 31 83 83 N to 52 N (Jereb et al., 2014) (Jereb et al., 2014) 52 83 N to 52 N (Jereb et al., 2014) (Jereb et al., 2014) 4 3 3 1 Inferred from video material in Golikov et al. 2023 (section (b) Behaviour (i) Feeding). Described as feeding behaviour using "spreading, enveloping and retreating" (see Discussion section) Golikov et al. (2023) 1 "spreading, enveloping and retreating" (Golikov et al. 2023) Golikov et al. (2023) 1 "spreading, enveloping and retreating" (Golikov et al. 2023) Golikov et al. (2023) NA 0 0 no ink sac in cirrates (Jereb et al., 2014) 0 Absent in cirrate octopuses (Jereb et al., 2014) (Jereb et al., 2014) 0 0 1 "beluga whales (Delphinapterus leucas)" (Choy et al 2016) Choy et al 2016 1 0 not gregarious (inferred from photo and video material) 1 0 1 0 "The eggs are laid singly on the
bottom and brooding duration is estimated to be about 20 to 32
months." (Jereb et al., 2014)		 Jereb et al., 2014 1 "The eggs are laid singly on the
bottom and brooding duration is estimated to be about 20 to 32
months." (Jereb et al., 2014)		 Jereb et al., 2014 8 7 12 75 NY
Cirrothauma murrayi Cirrothauma murrayi 4850 "Depth range from 2 430 to 4 850 m, with most captures
deeper than 3 000 m." (Jereb et al., 2014)		 (Jereb et al., 2014; Aldred et al 1983) 1000 "The cirrates mostly live below 1000 m, many in excess of 2000 m (Robson 1926, 1932
Voss 1967)." (Aldred et al 1983)		 (Jereb et al., 2014; Aldred et al 1983) 1 **edge case** "The photographs published by Roper & Brundage (1972) show cirrates to be suprabenthic, living close to the bottom but not actually in contact with it." (Aldred et al 1983). "Cirrothauma murrayi is a pelagic species that has been referred to as ‘the blind octopus’ because of its regressed eyes, although the eyes probably still function as photoreceptors (Aldred et al. 1983)." (Vecchione et al. 2010) Aldred et al 1983; Vecchione et al. 2010 61 85 40 N to 85 N (Jereb et al., 2014) (Jereb et al., 2014) 24 Bahamas "It is worth noting, however, that capture rate in the vicinity of the Blake Escarpment was twice as high as that of either the enclosed basins of the Bahamas
or the Middle Atlantic Continental Rise." (Vecchione 1987)		 (Vecchione 1987) 4 NA 0 2 2 1 "Also, they seem to mimic floating seaweed fragments and thus are difficult to detect (F. Gibran, pers. comm.)." (Anderson and Miriam 2020) Anderson and Miriam 2020 1 "It has also been seen using the arms and web as a "balloon" for defense, eliminating costly escape swimming (Boletzky et al., 1992)." (Seibel et al., 1997) Seibel et al., 1997 0 no ink sac in cirrates (Jereb et al., 2014) 0 Absent in cirrate octopuses (Jereb et al., 2014) (Jereb et al., 2014) 0 0 0 not gregarious (inferred from photo and video material) NA 0 NA "The small ‘light organs’ in the base of the sucker peduncles are present in both males and
females, and their function is presumably for mutual recognition for mating. The presence of light organs in other octopods has recently been demonstrated (Robison & Young 1981)." (Aldred et al 1983)		 "The small ‘light organs’ in the base of the sucker peduncles are present in both males and
females, and their function is presumably for mutual recognition for mating. The presence of light organs in other octopods has recently been demonstrated (Robison & Young 1981)." (Aldred et al 1983)		 9 7 8.2 155 NY
Cranchia scabra Cranchia scabra 2000 “Cranchia scabra has a very wide distribution.It has been captured from the surface to depths greater than 2000 m…” (Seibel et al., 1997). (Cairns, 1976; Jereb & Roper, 2010; Quetglas et al. 1999; Seibel et al., 1997) 0 "They are found distributed between 0 and 1500 m. There is no evidence of vertical
migration or ontogenic shift for this species" (Judkins & Vecchione, 2020)		 (Cairns, 1976; Jereb & Roper, 2010; Quetglas et al. 1999; Seibel et al., 1997) 1 "In the Gulf, C. scabra occupies a large section of the water column, from the surface down to 1500 m depth (Fig. 1D), with little evidence of ontogenetic shift or vertical migration (Judkins and Vecchione, 2020, in press)." (Timm et al., 2020) Timm et al., 2020 81 44 "C. scabra has been reported primarily from warm waters of the Atlantic, Pacific, and Indian oceans between about 35° N and 37 S (Clarke 1966). Pearcy (1965), however, recorded C. scabra from off Oregon at about 44° N. Seven VELERO specimens were taken from the Santa CataliNABasin at 33° N and one from zone 6 at 31°42' N." (Young, 1972) (Young, 1972) -37 "C. scabra is known from all temperate and tropical seas between 35 "N and 37' S (Clarke, 1966: 218)" (Cairns, 1976) (Cairns, 1976) 3 1 "The vertical distribution ranges from about 40 m to nearly 2 700 m with evidence of both significant ontogenetic descent as well as some diel vertical movement." (Jereb & Roper, 2010) (Jereb & Roper, 2010) 1 "The vertical distribution ranges from about 40 m to nearly 2 700 m with evidence of both significant ontogenetic descent as well as some diel vertical movement." (Jereb & Roper, 2010) (Jereb & Roper, 2010) 365.3 "a total lifespan of approximately 1 year (Arkhipkin 1996a; Arkhipkin and Nigmatullin, 1997)" (Arkhipkin, 2004:345) (Arkhipkin, 2004) 365.3 "a total lifespan of approximately 1 year (Arkhipkin 1996a; Arkhipkin and Nigmatullin, 1997)" (Arkhipkin, 2004:345) (Arkhipkin, 2004) Sexual maturity: The oldest immature individual of Arkhipkin's (1996) sample was 166 days. Unfortunately, no mature individuals were included in this study. "spend the first part of their lives (approximately 6 months) in tropical epi- and mesopelagic waters, and then descend to cold bathypelagic water to mature possibly for another 6 months" (Arkhipkin, 2004:345) (Arkhipkin, 2004:345) Sexual maturity: The oldest immature individual of Arkhipkin's (1996) sample was 166 days. Unfortunately, no mature individuals were included in this study. "spend the first part of their lives (approximately 6 months) in tropical epi- and mesopelagic waters, and then descend to cold bathypelagic water to mature possibly for another 6 months" (Arkhipkin, 2004:345) (Arkhipkin, 2004:345) NA 0 2 2 1 "Most cephalopods have a smooth skin so this roughness of the mantle in C. scabra may provide it with some protection as in the puffer fish. Indeed C. scabra can withdraw its tentacles, arms and head into the mantle cavity so that it resembles a ball, orange in colour when the chromatophores are expanded (Young, 1972; Angel, 1974). Small predators could be deterred by the transformation of the prey into a prickly ball; and the tubercles may give an illusion of greater size as well as breaking up the outline of the animal. It is known that C. scabra is eaten by some fish, Alepisaurus ferox (Rees & Maul, 1956) and tuNA(M. R. Clarke, personal communication) but both are large relative to the prey." (Dilly & Nixon 1976) Dilly & Nixon 1976
Nixon & Young 2003		 1 "The curious inking behaviour of the oceanic squid Teuthowenia (formerly Taonius) megalops may fall into the category of protean behaviour. Upon initial disturbance in a shipboard aquarium, small individuals will ink and jet away, but upon further disturbance they begin a ‘balling up’ sequence, in which the fin and eventually most of the mantle and head are inverted into the mantle cavity; the two tentacles are left extended but can be withdrawn too, and the animal will then ink inside the balled-up mantle (Dilly, 1972). R.E. Young (Reference Young1972b) noted that the oceanic squid Cranchia scabra reacts in much the same way." (Hanlon & Messenger, 2018, p. 130) Hanlon & Messenger, 2018, p. 130 1 (Roeleveld, 1977) "NA" (Xavier et al., 2018) 0 0 5 Found in the stomach of the blue shark (Prionace glauca) (Markaida & Sosa-Nishizaki, 2010) Markaida & Sosa-Nishizaki, 2010
Beasley et al. 2013
Antonelis et al. 1987
Imber, 1996		 3 0 not gregarious (inferred from photo and video material) NA 0 NA 19 14 138.8 125 NY
Discoteuthis laciniosa Discoteuthis laciniosa 1000 "Captured in open nets at depths between
100 and 1000 m." (Young & Roper, 1969)		 (Young & Roper, 1969; Nixon & Young 2003) 100 "Captured in open nets at depths between
100 and 1000 m." (Young & Roper, 1969)		 (Young & Roper, 1969; Nixon & Young 2003) 1 "Oceanic, mesopelagic " (Gomes-Pereira et al., 2016) Gomes-Pereira et al., 2016 19 33 From 33°04'N to 14°42'N (Young & Roper, 1969, p. 9) (Young & Roper, 1969, p. 9) 14 From 33°04'N to 14°42'N (Young & Roper, 1969, p. 9) (Young & Roper, 1969, p. 9) 2 NA 0 NA 0 1 Present in the family (Jereb & Roper, 2010) (Jereb & Roper, 2010) 0 0 2 bottlenose whales Hyperoodon planifrons off ArgentiNA(Clarke & Goodall 1994) Clarke & Goodall 1994
Clarke et al 1993		 1 0 not gregarious (inferred from photo and video material) NA 0 NA 1 1 97.6 51 NY
Egea inermis Egea inermis 2000 "The vertical distribution of paralarvae to approximately 35 mm mantle length is epipelagic, concentrated in subsurface water to 200 m both day and night. With growth and metamorphosis (around 35 to 40 mm mantle length) juveniles broaden their range in the mesopelagic zone to 800 m or more. One subadult female was captured in a closing net at night at '800 to 600 m in the western North Atlantic. Subadults and adults descend into the bathypelagic zone where maturation occurs in the 2 000 m zone. Captures of juveniles and large subadults at night in subsurface waters to about 300 m suggests that a portion of the population undergoes a diel vertical migration." (Jereb & Roper, 2010) (Cairns, 1976; Jereb & Roper, 2010) 300 [AL: No evidence of adult in epipelagic zone] "The vertical distribution of paralarvae to approximately 35 mm mantle length is epipelagic, concentrated in subsurface water to 200 m both day and night. With growth and metamorphosis (around 35 to 40 mm mantle length) juveniles broaden their range in the mesopelagic zone to 800 m or more. One subadult female was captured in a closing net at night at '800 to 600 m in the western North Atlantic. Subadults and adults descend into the bathypelagic zone where maturation occurs in the 2 000 m zone. Captures of juveniles and large subadults at night in subsurface waters to about 300 m suggests that a portion of the population undergoes a diel vertical migration." (Jereb & Roper, 2010) (Cairns, 1976; Jereb & Roper, 2010) 1 "The vertical distribution of paralarvae to approximately 35 mm mantle length is epipelagic, concentrated in subsurface water to 200 m both day and night. With growth and metamorphosis (around 35 to 40 mm mantle length) juveniles broaden their range in the mesopelagic zone to 800 m or more. One subadult female was captured in a closing net at night at '800 to 600 m in the western North Atlantic. Subadults and adults descend into the bathypelagic zone where maturation occurs in the 2 000 m zone. Captures of juveniles and large subadults at night in subsurface waters to about 300 m suggests that a portion of the population undergoes a diel vertical migration." (Jereb & Roper, 2010) Jereb & Roper, 2010 77 42 42 N to 35 S (Jereb & Roper, 2010) (Jereb & Roper, 2010) -35 42 N to 35 S (Jereb & Roper, 2010) (Jereb & Roper, 2010) 3 1 "The vertical distribution of paralarvae to approximately 35 mm mantle length is epipelagic, concentrated in subsurface water to 200 m both day and night. With growth and metamorphosis (around 35 to 40 mm mantle length) juveniles broaden their range in the mesopelagic zone to 800 m or more. One subadult female was captured in a closing net at night at 800 to 600 m in the western North Atlantic. Subadults and adults descend into the bathypelagic zone where maturation occurs in the 2 000 m zone." (Jereb & Roper, 2010) (Jereb & Roper, 2010) 0 "The vertical distribution of paralarvae to approximately 35 mm mantle length is epipelagic, concentrated in subsurface water to 200 m both day and night. With growth and metamorphosis (around 35 to 40 mm mantle length) juveniles broaden their range in the mesopelagic zone to 800 m or more. One subadult female was captured in a closing net at night at 800 to 600 m in the western North Atlantic. Subadults and adults descend into the bathypelagic zone where maturation occurs in the 2 000 m zone." (Jereb & Roper, 2010) (Jereb & Roper, 2010) NA 0 NA 0 0 0 0 not gregarious (inferred from photo and video material) NA 0 NA 5 4 53.2 198 NY
Eledone cirrhosa Eledone cirrhosa 800 Quetglas et al. (2000) found them from 50 m to 800 m depths (Salman et al., 2003; Belcari et al., 2002; Costa et al., 2005; Lauria et al., 2016; Quetglas et al., 2000; Salman et al., 2000; Sifner et al., 2005; Silva et al., 2011; Jereb et al., 2014; Sanchez & Martin 1993; Escánez et al. 2018; Gestal et al., 1999; Wurz et al. 1992; Pierce et al., 2010; Boyle, 1983; Jereb et al., 2015) 5 "Depths range from 5 to 500 m. Eledone cirrhosa occurs most commonly at depths between 60 to
120 m on the continental shelf throughout its range. " (Jereb et al., 2014)		 (Salman et al., 2003; Belcari et al., 2002; Costa et al., 2005; Lauria et al., 2016; Quetglas et al., 2000; Salman et al., 2000; Sifner et al., 2005; Silva et al., 2011; Jereb et al., 2014; Sanchez & Martin 1993; Escánez et al. 2018; Gestal et al., 1999; Wurz et al. 1992; Pierce et al., 2010; Boyle, 1983; Jereb et al., 2015) 2 "benthic as adults" (Salman et al., 2003) Salman et al., 2003 42 70 "Norwegian shelf up to 70°N" (Xavier et al., 2018) (Xavier et al., 2018) 28 "The occurrence of E. cirrhosa in the Canary Islands expands its current southern limit to about 28°N in the Atlantic Ocean." (Escánez et al. 2018) (Escánez et al. 2018) 1 1 “Distribution and abundance of Eledone cirrhosa, a benthic octopus inhabiting the NE Atlantic and Mediterranean Sea, were studied in north-western Iberian water. Capture data collected during two series of surveys, carried out during summer–autumn in Galician waters (NEAtlantic Ocean), were analyzed. ..Bathymetric distribution, variations in abundance, biomass and average body size of the octopus Eledone cirrhosa were analyzed through depth strata and seasons. Higher abundances were obtained in the intermediate strata and in autumn, most likely due to new recruits…Collected data and developed models illustrate a migratory behaviour during the reproductive period…, higher abundances were predicted near shore in early summer, corresponding to the beginning of spawning season. The models also indicate a displacement of higher abundance areas offshore through the summer, as suggested by Rees (1956). Late summer distribution models showed that the higher abundances were concentrated in deep waters (500–1000 m), which suggest a migration to deeper waters throughout the summer. Since our surveys did not include strata deeper than 500 m, the outputs of the models should be treated with caution“ (Regueira et al. 2014) (Regueira et al. 2014; Giordano et al. 2010; Relini et al. 2006; Wurtz et al. 1992; Elsevier, 2014) 1 "Although generally thought of as rather sedentary (Roper et al., 1984), evidence for a
pattern of seasonal migration in the Mediterranean is available from analyses of sex
ratio and maturity states from a range of depths. The deep-water population (100–200
m) normally has equal numbers of males and females, but trawls in shallower water
(60–90 m) in spring catch an increased number of maturing females. This seasonal sex
segregation is interpreted as a shoreward (shallower) migration of females for breeding
(Boyle, 1997). However, downward vertical migrations linked to spawning have also
been proposed. In Italian waters, there are more males than females at depths >300 m
(Belcari and Sartor, 1999a). In the Ligurian Sea, Orsi Relini et al. (2006), analysing 10
years of biological information, found a similar pattern and suggested that bathyal
hard substrata provide suitable seabed for egg laying and attachment. " (Jereb et al., 2015)		 (Villanueva & Norman, 2008) 1095.8 "In the western Mediterranean, lifespan appears to be 2 to 3 years and longer in colder waters further north in the eastern Atlantic" (Jereb et al., 2014) (Lefkaditou et al., 2001; Giordano et al. 2010; Relini et al. 2006; Jereb et al., 2015; Boyle & Rodhouse, 2005; Boyle, 1983; Boyle, 1983; Jereb et al., 2015; Pierce et al., 2010; Jereb et al., 2014; Regueira et al. 2015) 426.1 NW Iberia “Differential longevity between sexes was observed, with females reaching a life span of 17 months while males attained reached only 14 months, although the possibility that this difference might be even greater is discussed…Combining this with the results on age estimate obtained in this study (maximum age of 517 days), the life span of female E. cirrhosa should be around two years, agreeing with the hypothesis of Boyle (1986) for Scottish populations” (Regueira et al. 2015) (Lefkaditou et al., 2001; Giordano et al. 2010; Relini et al. 2006; Jereb et al., 2015; Boyle & Rodhouse, 2005; Boyle, 1983; Boyle, 1983; Jereb et al., 2015; Pierce et al., 2010; Jereb et al., 2014; Regueira et al. 2015) 700.1 Males 18-19 months and females 21-23 months with Mangold-Wirz
(1963) as reference in (Nixon, 1969)		 (Relini et al. 2006; Mangold-Wirz
(1963) as reference in (Nixon, 1969); Regueira et al. 2015)		 167 NW Iberia “Age of mature males ranged from 186 to 381 days (mean±SD: 292.8±74) on the northern fishing grounds, while it varied between 170 and 286 days (mean±SD: 243.7±40) in western waters. With the exception of one specimen with an estimated age of 167 days, the estimated age of mature females ranged between 310 and 453 days (mean±SD: 348.8±55) in the northern fishing grounds and from 386 to 450 days (mean±SD: 418±45) on the western fishing grounds (Fig. 5)….Based on size at first maturity (DML50%) estimated for this species on northwestern Iberian coasts (Regueira et al. 2013) and DML-Age relationships in this paper, females should achieve maturity at around one year old, while males should do so at approximately 8 or 9 months.” (Regueira et al. 2015) (Relini et al. 2006; Mangold-Wirz
(1963) as reference in (Nixon, 1969); Regueira et al. 2015)		 6 5 1 Hunting behaviour was found to show a consistent pattern. According to our observations, this behaviour is triggered by visually detection of its prey. When a prey is detected the octopus changes its chromatic pattern from a finely grained mottle of yellow-red colour with white spots (Fig. 1a) to an intense roughly grained reddish-colour, while gently approaches (Fig. 1b). When the octopus gets a distance of about one time its own mantle length (Fig. 1c), it pounces on and catches the crab using its brachial crown-umbrella complex (Fig. 1d), while maintaining the intense reddish coloured chromatic pattern" (Regueira et al. 2018) Regueira et al. 2018 1 Hunting behaviour was found to show a consistent pattern. According to our observations, this behaviour is triggered by visually detection of its prey. When a prey is detected the octopus changes its chromatic pattern from a finely grained mottle of yellow-red colour with white spots (Fig. 1a) to an intense roughly grained reddish-colour, while gently approaches (Fig. 1b). When the octopus gets a distance of about one time its own mantle length (Fig. 1c), it pounces on and catches the crab using its brachial crown-umbrella complex (Fig. 1d), while maintaining the intense reddish coloured chromatic pattern" (Regueira et al. 2018) Regueira et al. 2018
Regueira et al. 2018 (Lab)		 1 Hunting behaviour was found to show a consistent pattern. According to our observations, this behaviour is triggered by visually detection of its prey. When a prey is detected the octopus changes its chromatic pattern from a finely grained mottle of yellow-red colour with white spots (Fig. 1a) to an intense roughly grained reddish-colour, while gently approaches (Fig. 1b). When the octopus gets a distance of about one time its own mantle length (Fig. 1c), it pounces on and catches the crab using its brachial crown-umbrella complex (Fig. 1d), while maintaining the intense reddish coloured chromatic pattern" (Regueira et al. 2018) Regueira et al. 2018
(Hanlon and Messenger 1996; Mather et al. 2014)” (Regueira et al. 2018)		 1 It then positions its prey so that the anterior of the crab is close to its buccal mass (Grisley et al., 1996), and punctures it using its toothed radula and salivary papilla, which also has tooth-like protrusions at its tip (Nixon, 1979). This can be achieved either by boring a hole in the carapace (Boyle and Knobloch, 1981; Nixon & Boyle, 1982; Nixon and Maconnachie, 1988) or in the cornea of the crab (Grisley et al., 1996, 1999). It then injects saliva into this hole via the salivary papilla…crab. It is paralyzed in 1-2 min, and respiration is strongly inhibited (Ghiretti, 1960; Boyle, 1990), although the heart it still beating at this point, and dies about 10 min later. The beak and radula are used to remove the flesh from the crab carapace.” (Key et al. 2002) Key et al. 2002
Runham et al. 1997
(Nixon & Boyle 1982)
Hiemstra, 2015
Matthews-Cascon et al., 2009
(Boyle & Knobloch, 1981; Nixon & Boyle, 1982)" (Guerra & Nixon, 1987)		 1 observations from aquarium “an octopus will move stealthily towards a crab and envelop it in its web of suckers to ensure rapid immobilizations. (Key et al. 2002) (Key et al. 2002) 1 "The hunting behaviour observed in E. cirrhosa has one feature similar to "stalking" because the octopus "jumps" on the prey. However, a true stalking involves a head-first attack after a long tracking phase, in which the predator gradually closes on the prey from behind, where its visibility is poorest (Hanlon and Messenger 1996). These two phases of this hunting tactic were not observed in the horned octopus. On the other hand, stalking seems to be reserved for large, fast prey, while slower fish are caught by the squids without stalking, as occurred in E. cirrhosa" (Regueira et al. 2018) Regueira et al. 2018
Grisley et al. 1999		 2 1 1 “‘Sand covering’ similar to the behaviour seen in Sepia spp. and various species of sepiolid, was described for the first time in the horned octopus by Guerra et al. (2006). The animals displayed a burying behaviour lasting ca. 50 s. The behaviour continued until the animal was totally covered except for its eyes” (Regueira et al. 2018) Regueira et al. 2018
Guerra et al. 2006
Jereb et al., 2015		 1 (Jereb et al., 2014) 39 "The diet of E. cirrhosa, has been described as consisting mainly of crustaceans, but fish, cephalopods, polychaetes and gastropods have also been reported (Sánchez 1981; Boyle 1983; Grisley et al. 1999) and evidence of bivalve prey was noted once by Boyle (1986)." (Costa et al., 2005) Costa et al 2005
Regueira et al. 2017
Jereb et al., 2014
Xavier et al 2018
Relini et al. 2006
Pierce et al., 2010		 0 0 6 27 "was the single most important prey species of Tope (Galeorhinus galeus (Linnaeus, 1758)) and the second most important in Greater Spotted Dogfish (S. stellaris (Linnaeus, 1758)) (Ellis et al. 1996). E. cirrhosa beaks were also found in the stomach of a juvenile six-gill shark (Hexanchus griseus (Bonnaterre, 1788)) captured off Inishbofin Island, Co. Galway during September 1986 (Quigley and Flannery unpublished data, NMINH:1991.43.1)." (Quigley & Flannery 2014) Jereb et al 2015
Quigley & Flannery 2014		 5 2 seem relatively tolerant in lab conditions. mature/spawning females in lab "neither territorial behaviour nor hostility was observed among the animals sharing the same tank during this study. This fact could be due to the similar sizes of the specimens sharing each tank as well as keeping the animals fed at libitum" (Regueira et al. 2018) Regueira et al. 2018 0 seem relatively tolerant in lab conditions. mature/spawning females in lab "neither territorial behaviour nor hostility was observed among the animals sharing the same tank during this study. This fact could be due to the similar sizes of the specimens sharing each tank as well as keeping the animals fed at libitum" (Regueira et al. 2018) Regueira et al. 2018 2 1 1 "Eleven octopi (Eledone cirrhosai were deprived of food for 24 h and then were presented individually a paper model of a crab, a live crab as a model, and a neutral letter T on the outside of the testing tank for 5,10, and 3 days, respectively. When there was no approach to the model
for 20 min, the model was removed and a live crab was introduced into the tank. The results
indicated that (1) octopi are capable of inhibitory learning, (2)they can transfer the learned inhibition to a natural feeding situation, (3)they remember the learned inhibitions for at least 37
days"		 (ANGERMEIER & Dassler 1992) not specifically mentioned in ethogram Boyle & Dubas 1980 1 0 lab "The first spawning female, placed in the individual tank, performed a burrowing behaviour, which can be summarized as follows: i) the spawning female always remained inside the shelter; ii) she blocked the entrance with stones and/or other different materials available from the adjacent environment (Fig. 3), iii) the animal showed an increasingly physical degradation (e.g. skin ulcerative lesions, low ingestion rate, if any, etc.) during the "incubation" time"…our observations confirm that horned octopus females perform an active care of the egg lying until their death, as many other benthic cephalopod species do (Hanlon and Messenger 1996)." (Regueira et al. 2018) (Regueira et al. 2018) 1 lab "The first spawning female, placed in the individual tank, performed a burrowing behaviour, which can be summarized as follows: i) the spawning female always remained inside the shelter; ii) she blocked the entrance with stones and/or other different materials available from the adjacent environment (Fig. 3), iii) the animal showed an increasingly physical degradation (e.g. skin ulcerative lesions, low ingestion rate, if any, etc.) during the "incubation" time"…our observations confirm that horned octopus females perform an active care of the egg lying until their death, as many other benthic cephalopod species do (Hanlon and Messenger 1996)." (Regueira et al. 2018) (Regueira et al. 2018) 230 182 394.68 187.44 110 66 W-W
Eledone moschata Eledone moschata 612 "It is mainly distributed at
depths of 15–200 m in both Mediterranean waters and Iberian waters of the Gulf of
Cádiz, where it is most abundant in shallow waters down to 100 m (Gamulin-Brida
and Ilijanić, 1972; Salman et al., 1997; Lefkaditou et al., 1998a; Belcari et al., 2002a; Silva
et al., 2004). In the northern Adriatic, densities were nearly 700 km –2 at 10–50 m, but
decreased to <300 km –2 at 50–100 m and to ca. 30 km –2 at 100–200 m (Krstulović Šifner
et al., 2011). In some areas, it is found at greater depths: to 450 m in the Gulf of Cádiz
(Silva et al., 2004), 612 m in southern Portuguese waters (Lourenço et al., 2008), and 320 m in the Aegean Sea (Salman et al., 2000)." (Jereb et al., 2015)		 (Jereb et al., 2014; Akyol et al., 2007; Belcari et al., 2002; Costa et al., 2005; Krstulocic Sifner & Vrgoc, 2009; Lauria et al., 2016; Oellermann et al., 2015; Quetglas et al., 2000; Salman et al., 2000; Sen & Akyol, 2011; Sifner et al., 2005; Sifner & Vrgoc, 2009; Silva et al., 2004; Silva et al., 2011; Escánez et al. 2018; Ruby and Knudsen 1972; Adam, 1937; Pierce et al., 2010; Jereb et al., 2015; Boyle, 1983; Boyle, 1983; Mather et al., 2012) 0 " occurs all over the Mediterranean Sea mainly at depths between 0 – 200 m" (Oellermann et al., 2015) (Jereb et al., 2014; Akyol et al., 2007; Belcari et al., 2002; Costa et al., 2005; Krstulocic Sifner & Vrgoc, 2009; Lauria et al., 2016; Oellermann et al., 2015; Quetglas et al., 2000; Salman et al., 2000; Sen & Akyol, 2011; Sifner et al., 2005; Sifner & Vrgoc, 2009; Silva et al., 2004; Silva et al., 2011; Escánez et al. 2018; Ruby and Knudsen 1972; Adam, 1937; Pierce et al., 2010; Jereb et al., 2015; Boyle, 1983; Boyle, 1983; Mather et al., 2012) 2 "This coastal species occurs on muddy substrates." (Jereb et al., 2014) Belcari et al., 2002;
Jereb et al., 2014;
Lauria et al., 2016;
Sen & Akyol, 2011; 		18 46 46 N to 30 N (Jereb et al., 2014) (Jereb et al., 2014) 28 "the distribution range of E. moschata is extended here [Canary Islands] from 40°N to 28°N in the eastern Atlantic." (Escánez et al. 2018) (Escánez et al. 2018) 1 1 "Mangold-Wirz (1963) and Mangold (1983) stated that the reproductive migrations affected the sex ratio in relation to size in particular areas and seasons. Furthermore, Ezzedine-Najai (1997) also indicated that the mature females leave the fishing grounds during the spawning season, hence favouring the predominance of males in the sex ratio. But Silva et al. (2004) reported that this phenomenon was not observed, and contrarily, there were a major proportion of females observed throughout the sample period. In this study, except in December and January, females were in excessive proportion, the same as the findings of Silva et al. (2004). Only during the January, spawning period for females (male:female, 1:0.54), there was no evidence of mature females leaving the fishing grounds." (Akyol et al., 2007) (Akyol et al., 2007; Krstulocic Sifner & Vrgoc, 2009; Jereb et al., 2015; Boyle, 1983) 1 "In February and March the individuals congregate in shallow water. The large animals
disappear again in May after breeding; only the young non-breeding individuals remain,
and do not migrate to deeper water until November- December" (Ruby and Knudsen 1972)		 (Ruby and Knudsen 1972; Salman et al., 2000; Boyle, 1983) 730.5 "Lifespan is probably up to 2 years (e.g.
Mangold, 1983b)." (Jereb et al., 2015)		 (Jereb et al., 2015; Boyle, 1983) 456.6 "The entire life-span for females. including the long brooding period. would vary between 15 and probably 23 months" (Boyle, 1983). (Jereb et al., 2015; Boyle, 1983) 700.1 Males 15-16 months and females 21-23 months with Mangold-Wirz
(1963) as reference in (Nixon, 1969)		 (Boyle, 1983; Mangold-Wirz
(1963) as reference in (Nixon, 1969))		 273.9 "Laboratory-reared females of Eledone moschata spawn at the age of 9 to 12 months" (Boyle, 1983). (Boyle, 1983; Mangold-Wirz
(1963) as reference in (Nixon, 1969))		 4 4 1 "(Lab Study) Octopuses usually moved on the bottom of the tank with arms extended in all directions, and with the web tilted upward in the direction of progress. The two leading arms were often held off the bottom and extended into the water in front of the octopus, a posture which would trap prey such as crabs." (Mather, 1985) (Mather, 1985) 1 "Boletzky (1977) observed them chase and attack newly-hatched Sepia officianalis the bottom" (Boyle, 1983). Boletzky 1977
Boyle, 1983		 1 "E. moschata caught crabs by surrounding them with the arm web and holding them with the suckers on their arms. […] In 10 cases the octopus turned toward the crab and extended an arm tip toward it, only holding it with all arms after initial contact was made. in eight cases the octopus only captured the crab after tactile contact with an arm or web (sometimes initiated by the crab). (Mather 1985) Mather 1985 1 On the other hand, it was rare to find exoskeletons of crustaceans in stomachs of larger specimens (ML . 80 mm): these usually contained only highly macerated soft parts of the prey’s body, suggesting that larger individuals drilled holes in the exoskeletons of their preys and sucked out the soft parts (or they just removed the flesh from the exoskeleton to eat it). It was previously reported that juvenile and adult E. moschata drilled the carapaces of crustaceans (Boletzky, 1975). In aquarium bred E. moschata it was observed that adult animals fed by drilling holes in carapaces of crustaceans more often thanjuveniles, and in the related species. (Krstulocic Sifner & Vrgoc, 2009) Krstulocic Sifner & Vrgoc, 2009 2 1 1 "They are able to hide. burying themselves in soft bottom (Boletzky. 1975d). Boletzky. 1975d 1 (Jereb et al., 2014) 24 "The major prey categories found in stomachs of the musky octopus with the highest percentage of occurrence (%F) were crustaceans (65.0%), fish (37.8%) and cephalopods (21.8%). The most abundant considering number (%N) were crustaceans (57.1%), followed by fish (28.0%), cephalopods (10.8%), gastropods (2.1%), polychaetes (1.3%) and bivalves (0.6%). Crustaceans were also most important considering weight (40.1%), followed by fish (31.5%) and cephalopods (23.7%). Polychaetes and gastropods were found with weight percentage 2.6% and 1.9%, respectively (Figure 2). The rest of the prey appeared in stomachs only sporadically, accounting for less than 0.5% in weight and number, and they were not considered as prey groups (species belonging to Pteropoda, Porifera, Foraminifera, Trematoda and Xantophyceae)." (Krstulocic Sifner & Vrgoc, 2009) Jereb et al., 2014
Costa et al., 2005
Krstulocic Sifner & Vrgoc, 2009
Sen & Akyl, 2011		 0 1 Krstulocic Sifner & Vrgoc, 2009
Jereb et al., 2015		 6 9 Found in the stomach of juveniles and maturing Mustelus punctulatus (Di Lorenzo et al., 2020) Di Lorenzo et al., 2020
Jereb et al 2015		 5 2 "Yet Mangold-Wirz (1959) reports that Eledone moschata were found clustered in the Mediterranean Sea, a pattern that could either denote social grouping, or spacing in response to a patchy distribution of food or shelter. " (Mather, 1985) Mather 1985 0 "Yet Mangold-Wirz (1959) reports that Eledone moschata were found clustered in the Mediterranean Sea, a pattern that could either denote social grouping, or spacing in response to a patchy distribution of food or shelter. " (Mather, 1985) Mather 1985 4 2 1 Marini et al. 2017 1 Marini et al. 2017 2 1 "Nine encounters were designated as mating interactions, since a male either attempted to insert, or succeeded in inserting, the third right arm (hectocotylus) into the mantle cavity of a female. In seven of these attempts the male used the web-spread posture over the female, but this posture was maintained for the whole period in only three encounters. No special colour pattern accompanied mating, which occurred at several times of day. Females were observed to refuse mating by closing the mantle cavity and/or pushing the male's arm away. Mating ended when the female pushed the male away (five occurrences), when another male interfered (three occurrences), or when the male left (three occurrences). In some cases mating was resumed later. The longest duration was 18 rain. The mating process, despite the web-spread clasp by the male, was not accompanied by fighting. " (Mather, 1985) (Mather, 1985) 0 "Nine encounters were designated as mating interactions, since a male either attempted to insert, or succeeded in inserting, the third right arm (hectocotylus) into the mantle cavity of a female. In seven of these attempts the male used the web-spread posture over the female, but this posture was maintained for the whole period in only three encounters. No special colour pattern accompanied mating, which occurred at several times of day. Females were observed to refuse mating by closing the mantle cavity and/or pushing the male's arm away. Mating ended when the female pushed the male away (five occurrences), when another male interfered (three occurrences), or when the male left (three occurrences). In some cases mating was resumed later. The longest duration was 18 rain. The mating process, despite the web-spread clasp by the male, was not accompanied by fighting. " (Mather, 1985) (Mather, 1985) 0 "Boletzky (1975) reared Eledone moschata from hatching to a maximal size of 4 10 g in 10.5 months. Spawning occurred at this time and females lived another two months while brooding the eggs." (Forsythe 1984) (Forsythe 1984) 1 "Boletzky (1975) reared Eledone moschata from hatching to a maximal size of 4 10 g in 10.5 months. Spawning occurred at this time and females lived another two months while brooding the eggs." (Forsythe 1984) (Forsythe 1984) 220 73 276.336 150.252 59.2 107 66 W-W-NY
Enteroctopus dofleini Enteroctopus dofleini 1500 "Depths range from 0 to 1 500 m." (Jereb et al., 2014) (Villanueva & Norman, 2008; Jereb et al., 2014; AZA Aquatic Invertebrate Taxon Advisory Group, 2014; Hollenbeck & Scheel, 2017; Conrath & Sisson, 2018; Brewer et al., 2017; Yoon et al., 2016; Scheel, 2015; Sano & Bando, 2015; Toussaint et al., 2012; Scheel, 2002; Ikeda et al., 1999; Kubodera, 1991; Boyle, 1983; 0 "Depths range from 0 to 1 500 m." (Jereb et al., 2014) (Villanueva & Norman, 2008; Jereb et al., 2014; AZA Aquatic Invertebrate Taxon Advisory Group, 2014; Hollenbeck & Scheel, 2017; Conrath & Sisson, 2018; Brewer et al., 2017; Yoon et al., 2016; Scheel, 2015; Sano & Bando, 2015; Toussaint et al., 2012; Scheel, 2002; Ikeda et al., 1999; Kubodera, 1991; Boyle, 1983; 2 "typically occurs in dens on rocky reefs or boulder areas with sand-shell substrate; the same den is utilized for up to several months. Individuals also have been observed in sand and mud habitats. At the northern end of its range, it occurs commonly on reefs in the intertidal zone" (Jereb et al., 2014) Jereb et al., 2014
Heery et al., 2018
Scheel, 2002		 36 66 66 N to 30 N (Jereb et al., 2014) (Jereb et al., 2014) 30 66 N to 30 N (Jereb et al., 2014) (Jereb et al., 2014) 1 1 "dofleini migrates offshore into deeper water in summer in order to mate. In autumn and
winter animals migrate back inshore where eggs are spawned" (Jereb et al., 2014)		 (Jereb et al., 2014; Hollenbeck & Scheel, 2017; Brewer et al., 2017; Scheel, 2015; Larson et al., 2015; Toussaint et al., 2012; Scheel & Bisson, 2012; Boyle, 1983; Elsevier, 2014;) 1 "dofleini migrates offshore into deeper water in summer in order to mate. In autumn and
winter animals migrate back inshore where eggs are spawned" (Jereb et al., 2014)		 (Jereb et al., 2014; Villanueva & Norman, 2008; Sano & Bando, 2018; Scheel, 2015; Boyle, 1983; ) 1826.3 3-5 years (Nafkha et al., 2019) (Villanueva & Norman, 2008; Anderson et al., 2002; AZA Aquatic Invertebrate Taxon Advisory Group, 2014; Hollenbeck & Scheel, 2017; Hartwick, 1983 in Boyle, ed.; Brewer et al., 2017; Boyle, 1983; Hanlon and Messnger 2018 – Book p 242; Boyle & Rodhouse, 2005; Conrath & Conners, 2014; Hartwick, in Boyle (1983). p. 286;Nafkha et al., 2019; Iglesias et al., 2014) 1095.8 3-5 years (Nafkha et al., 2019) (Villanueva & Norman, 2008; Anderson et al., 2002; AZA Aquatic Invertebrate Taxon Advisory Group, 2014; Hollenbeck & Scheel, 2017; Hartwick, 1983 in Boyle, ed.; Brewer et al., 2017; Boyle, 1983; Hanlon and Messnger 2018 – Book p 242; Boyle & Rodhouse, 2005; Conrath & Conners, 2014; Hartwick, in Boyle (1983). p. 286;Nafkha et al., 2019; Iglesias et al., 2014) 1095.8 "Martin (pers. comm) believes that the average time to maturity is 3 years but that males may live 1 or 2 years longer if they don't reproduce" (Hartwick, i Boyle (1983). p. 286) (Hartwick, i Boyle (1983). p. 286; Boyle, 1983; Boyle & Rodhouse, 2005; Hartwick, 1983 in Boyle, ed.) 730.5 Sexual maturity: 2-3 years (Hartwick, 1983 in Boyle, ed.) (Hartwick, i Boyle (1983). p. 286; Boyle, 1983; Boyle & Rodhouse, 2005; Hartwick, 1983 in Boyle, ed.) 7 6 1 "hunts mostly by speculation using the tips of its arms to probe around and under rocks (personal observation)" (Corsgove, 2002) Cosgrove, 2002
Hanlon & Messenger, 2018, p. 84		 1 "In most cases the prey is either captured or flushed out, after which the pursuit mode of hunting is employed. When prey is discovered in the open the pursuit mode is also employed." (Cosgrove, 2002) (Cosgrove, 2002) 1 Johnson (1942) noted that Enteroctopus dofleini would ambush by “suddenly striking out at prey with a tentacle and grasping it with the aid of the suckers”, and he noted that the octopus was ineffective using this method while ambushing shrimp and small fishes that lived among seaweeds. (Cosgrove 2002) Cosgrove 2002 1 "the webover technique" (Cosgrove, 2002) Cosgrove, 2002
Mather (1991, 1998)		 1 – Toxic bite (Songdahl & Shapiro, 1974)
– "Shelled prey are drilled" (Jereb et al., 2014)		 Songdahl & Shapiro, 1974
Mather, 1998
Jereb et al., 2014
Hiemstra, 2015
Anderson et al., 2008
AZA Aquatic Invertebrate Taxon Advisory Group, 2014
Ambrose et al., 1988
Matthews-Cascon et al., 2009		 1 " Suckers located on their arms are able to create tremendous force in prying apart shells, and their arms are limber and long enough to reach the most elusive prey." (AZA Aquatic Invertebrate Taxon Advisory Group, 2014) AZA Aquatic Invertebrate Taxon Advisory Group, 2014 1 – "the use of the net is an example of the adaptability in foraging behaviour suggested in other studies of octopus behaviour." (Rigby & Sakurai, 2005)
– "[T]he remarkable net-climbing behaviour of
Enteroctopus dofleini [formerly Octopus dofleini) provides
further evidence of the flexibility of cephalopod
behaviour, in foraging as in other activities.
A radio acoustic tagging study (Chapter 1] of the Giant Pacific
Octopus in Northern Japan, combined with SCUBA
diving, has shown that this species can move from a den
towards a commercially deployed gill net, climb it, and
feed on the ensnared fish, before descending and
returning to the den (Rigby 8: Sakurai, 2005)." (Hanlon & Messenger, 2018, p. 89)
– "E. dofleini has primarily been reported as being nocturnal (Mather et al., 1985; Scheel and Bisson, 2012), but can modify their behavior based on foraging opportunities (Rigby and Sakurai, 2005)." (Hofmeister & Voss 2017)		 Rigby & Sakurai, 2005
Hanlon & Messenger, 2018, p. 89
Hofmeister & Voss 2017
Anderson and Mather (2007)
Onthank & Cowles, 2011
Anderson & Mather, 2007
Hartwick et al., 1978
Hanlon & Messenger, 2018, p. 94		 6 6 1 Den use (Scheel, 2002) Scheel, 2002
Mather et al. (1985) in Hofmeister & Voss, 2017
Jereb et al 2014		 1 "Burrowing is performed only by octopuses and occurs in a variety of habitats. For example, Octopus dofleini burrows in sand, or sand and mud mixtures (Hartwick, Thorarinsson & Tulloch, Reference Hartwick, Thorarinsson and Tulloch1978 " (Hanlon & Messenger, 2018, p. 121) Hanlon & Messenger, 2018, p. 121 1 "Scheel et al. (2007) studying E. dofleini in Alaska found that octopuses were selective of prey species and size in a manner consistent with a rate-maximizing optimal forager, while the unrelatedness of octopus and prey population trends was consistent with a risk-minimizing forager. They hypothesized that octopuses may rate-maximize while foraging and act as a risk-minimizing forager by decreasing movement between foraging patches." (Onthank & Cowles, 2011) Onthank & Cowles, 2011 1 Sucker display is mentioned as a defensive posture (Anderson et al., 2010) Anderson et al., 2010 1 "Functional ink ejection has been observed in captive hatchlings, such as Enteroctopus dofleini (yamashita 1974, Gabe 1975, okubo 1979)" (Villanueva & Norman, 2008) Villanueva & Norman, 2008
AZA Aquatic Invertebrate Taxon Advisory Group, 2014		 1 "E. dofleini will normally move under cover or will flee to deeper water (personal observation" (Cosgrove, 2002) Cosgrove, 2002 64 "Diet consists mainly of bivalve and gastropod molluscs and decapod crustaceans. Shelled prey are drilled. Other prey includes echinoderms, brachiopods, assorted fishes, shark eggs, and even seabirds." (Jereb et al., 2014) Jereb et al 2014
Xavier et al., 2018
Heery et al., 2018
Scheel & Anderson 2012
Sano et al., 2017
Anderson & Shimek, 2014		 1 Heery et al., 2018 1 Kanamaru (1964) in (Villanueva & Norman, 2008) 8 12 Found in stomachs of Steller sea lions (Eumetopias jubatus) (Sinclair, 2019) Antonelis et al. 1987
AZA Aquatic Invertebrate Taxon Advisory Group, 2014
Sinclair, 2019
Hartwick & Thorarinsson, 1978
Xavier et al 2018		 4 1 "Mather, Resler & Cosgrove (unpublished), when tracking O. dofleini with sonic tags, found that individuals seldom came within 100 m of each other. " (Mather, 1985) AZA Aquatic Invertebrate Taxon Advisory Group, 2014; Mather, 1985 0 "Mather, Resler & Cosgrove (unpublished), when tracking O. dofleini with sonic tags, found that individuals seldom came within 100 m of each other. " (Mather, 1985) AZA Aquatic Invertebrate Taxon Advisory Group, 2014; Mather, 1985 12 10 1 Habituation to objects (bottles of varying colours and textures) demonstated in trials “Every octopus habituated to the object’s presence within trials, in that the criterion of 30 min without contact was met during every trial; contact time in trials ranged from 0 to 127 minutes…Baldwin and Baldwin’s (1997) comments on the slower habituation expected to a more complex stimulus situation is appropriate here” Mather & Anderson 1999 1 Marini et al. 2017 1 “Octopuses followed bathymetry contours on central-tendency movements, resulting in loops and a return to a den, while direct movements crossed bathymetry contours, and resulted in relocation (Fig. 5). In Prince William Sound, bare rock is exposed as vertical cliffs and octopuses may be found crawling along the bottom of cliff edges or hiding in crevices or dens along cliff faces. Thus, visually and tactually conspicuous edges run parallel to bathymetry contours in rocky habitats where octopuses are found and these intermittent tracking samples were obtained. We interpret these movement patterns as reliance on local submarine topography for navigation…Central-tendency movements recorded by intermittent tracking were oriented parallel to contours, while movements without central-tendency crossed contours, suggesting that animals navigate by contour following to return to a known den." (Scheel & Bisson, 2012) 1 "Visual discrimination training has also been
successfully carried out with 0. apollyofli 0- may“-
O. bimaculatus and O. bimaculoides (Roffe, 1975;
Hanlon, Forsythe & Messenger, 1984; Allen, Michels
& Young, 1986; Boal, 1991], and with the decapods Sepia
officinalis (Messenger, l977b), Lolliguncula brevis
(Allen, Michels st Young, teas) and Todarodes P“°lfif"$
(Flores, 1983)."		 Hanlon and Messenger 2018 : 229 1 “Octopus adaptability has been used for problem solving across domains. For example, octopuses removed a lid of a glass jar to obtain a crab confined inside, but Fiorito, von Planta and Scotto (1990) found that they did not learn (i.e., did not have a shorter latency to perform the task). Anderson & Mather (2010) added an extra cue, chemical traces on the outside of the jar, which did produce a decrease in latency. Because the jar was commonly taken under the arm web and out of sight, the octopus did not remember the situation without continuing cueing, but could use the two cues sequentially. There was no stereotyped behavior in this novel situation; one individual tested by the author simply wrapped one arm around the edge of the lid and contracted it to unscrew the lid; the other used the bases of several arms and moved them laterally." (Mather, 2019) Mather, 2019 1 “Octopuses hold a mollusk or other prey item within the grasp of the proximal arms clearly out of their sight, so they cannot be using vision to mediate this set of behaviors. Because they can realize the failure of pulling and switch to another technique (Fiorito & Gherardi, 1999), as well as manipulate the position of shelled mollusks so that they can choose the most effective position to drill or chip, based on past learning or new evidence, tactile–proprioceptive information about arm position is being stored and used.” (Anderson & Mather 2007) (Anderson & Mather 2007) 1 long-term (>1 days) recognition of individual humans “These exposures were given once in the morning and once in the afternoon at least 4 hr apart for 5 days, followed by a 2-day break with no feeding and no human interaction; the treatment was repeated for 5 more days. After this regimen, in a follow-up on the 11th day, each person just looked in the tank and recorded octopus reactions, 10 min apart, reversing the order of human presentation with each octopus…By the end of testing, four responses stood out as differing between feeders and irritators: direction of movement, a dark Eyebar body pattern, direction of funnel aiming, and respiration period.” (Anderson et al. 2010) Anderson et al. 2010 1 "Mather & Anderson (1999) set up a ‘boring’ laboratory situation to elicit play behavior
from E. dofleini. Isolated octopuses were given a floating pill bottle. Their initial reaction was
to grasp it and bring it under the arm web to the mouth. By the third of ten trials, the
octopuses ignored it, but by the fifth or sixth trial, two individuals began to aim jets of water
at the bottle, which sent it to the other end of the tank, only to have it returned by the current
of incoming water — and the octopuses jetted at it again, the marine equivalent of bouncing
a ball. Based on Burghardt’s (2005) definition, these actions were play."		 (Mather, 2019) 1 use of water as tool "Mather & Anderson (1999) set up a ‘boring’ laboratory situation to elicit play behavior from E. dofleini. Isolated octopuses were given a floating pill bottle. Their initial reaction was to grasp it and bring it under the arm web to the mouth. By the third of ten trials, the octopuses ignored it, but by the fifth or sixth trial, two individuals began to aim jets of water at the bottle, which sent it to the other end of the tank, only to have it returned by the current of incoming water — and the octopuses jetted at it again, the marine equivalent of bouncing a ball. Based on Burghardt’s (2005) definition, these actions were play." (Mather, 2019) 1 "Because they can realize the failure of pulling and switch to another technique
(Fiorito & Gherardi, 1999), as well as manipulate the position
of shelled mollusks so that they can choose the most effective
position to drill or chip, based on past learning or new evidence, tactile–proprioceptive information about arm position is being
stored and used. Such a result, that the octopus is performing trial-and-error
learning and switching penetration techniques when the initial one
fails to work (Fiorito & Gherardi, 1999; Wodinsky, 1973)" (Anderson & Mather, 2007); "Anderson & Mather (2010) added an extra cue, chemical traces on the outside of the jar, which did produce a decrease in latency. Because the jar was commonly taken under
the arm web and out of sight, the octopus did not remember the situation without continuing
cueing, but could use the two cues sequentially. There was no stereotyped behavior in this
novel situation; one individual tested by the author simply wrapped one arm around the edge
of the lid and contracted it to unscrew the lid; the other used the bases of several arms and
moved them laterally." (Mather, 2019)		 Anderson & Mather 2007; Mather 2019 2 0 "Like most octopuses, Enteroctopus dofleini excavates dens in suitable substrate, and associated with this octopus were scavenging fish, sea stars and crustaceans (Hartwick and Thorarinsson 1978). Active predators, particularly groupers, also associate with tropical foraging octopuses and thereby obtain access to otherwise inaccessible prey (Diamant and Shpigel 1985)…Hartwick and Thorarinsson (1978) also reported fish, primarily various sculpins, that cohabitated within the dens of Enteroctopus dofleini." (Scheel et al. 2014) (Scheel et al. 2014) 1 "Sophisticated pre-copulatory behaviors by the male are not
uncommon, and include elaborate changes in coloration and aggressive movements towards potential
competitors." (AZA Aquatic Invertebrate Taxon Advisory Group, 2014)		 (AZA Aquatic Invertebrate Taxon Advisory Group, 2014) 0 "Several female GPOs have been reported to continue to eat during the first
part of egg guarding" (Anderson et al., 2002)		 (Anderson et al., 2002) 1 "Several female GPOs have been reported to continue to eat during the first
part of egg guarding" (Anderson et al., 2002)		 (Anderson et al., 2002) 205 115 1338.8 NY
Galiteuthis glacialis Galiteuthis armata 2500 "The vertical distribution of paralarvae and juveniles extends from the lower epipelagic zone, concentrated around 200 m, but as shallow as around 75 m. With continued growth ontogentic descent finds larger juveniles concentrated at 300 to 1 000 m, subadults to 2 000 m; then sexual maturation occurs in very deep water in excess of 2 500 m. Significant diel vertical shifting appears to occur. " (Jereb & Roper, 2010) (Jereb & Roper, 2010; Sajikumar et al. 2020; Piatkowski & Hagen 1994; Roper 1969; Roper & Young 1975; 500 [AL: paralarvae higher but no evidence of non post-spawning moribund adults above 500 m] "The vertical distribution of paralarvae and juveniles extends from the lower epipelagic zone, concentrated around 200 m, but as shallow as around 75 m. With continued growth ontogentic descent finds larger juveniles concentrated at 300 to 1 000 m, subadults to 2 000 m; then sexual maturation occurs in very deep water in excess of 2 500 m. Significant diel vertical shifting appears to occur. " (Jereb & Roper, 2010) paralarvae caught at 500m in Antarctic waters. "Nesis (1998) hypothesized that G. glacialis spawning occurs at the same depth of the adult habitat (500–2500 m)" (Sajikumar et al. 2020) "As with other cranchiids, adults occur in the lower mesopelagic and bathypelagic zones (Nesis et al. 1998), whereas post-spawning moribund squids become positively buoyant and appear near the surface, where they fall prey to seabirds (Croxall & Prince 1996)." (Laptikhovsky & Arkhipkin 2003) (Jereb & Roper, 2010; Sajikumar et al. 2020; Piatkowski & Hagen 1994; Roper 1969; Roper & Young 1975; 1 in Antarctic "The species was mainly concentrated in the open ocean as well as along the steep continental slope of the eastern Weddell Sea. G. glacialis did not occur in shallow shelf regions or the southern Weddell Sea, but preferred the oceanic regions, which is typical for all cranchiid squids (Voss 1980, Nesis 1987)." (Piatkowski & Hagen 1994) Piatkowski & Hagen 1994 26 -44 "our records are the first outside circumpolar Antarctic waters, thus expanding the geographical distribution of the species towards the north, at least until 44°02'95" South. SST [sea surface temperature] and SBT [sea bottom temperature] seem to indicate that this species was caught along the western border of the FMC [Falkland
(Malvinas) Current]" (Guerra et al. 2011)		 (Guerra et al. 2011) -70 "45°S to 70°S" (Jereb & Roper, 2010) (Jereb & Roper, 2010) 3 1 "The ASC [Antarctic Slope Current] which has slow movement with an average speed of 8 cm s−1 (Thompson et al., 2018) may result in the paralarvae and juveniles being carried westward from the spawning area on the continental shelf edge near Prydz Bay." (Sajikumar et al. 2020) (Sajikumar et al. 2020) 1 "The vertical distribution of paralarvae and juveniles extends from the lower epipelagic zone, concentrated around 200 m, but as shallow as around 75 m. With continued growth ontogentic descent finds larger juveniles concentrated at 300 to 1 000 m, subadults to 2 000 m; then sexual maturation occurs in very deep water in excess of 2 500 m. Significant diel vertical shifting appears to occur. " (Jereb & Roper, 2010) (Roper & Young 1975; Jereb & Roper, 2010; Piatkowski & Hagen 1994) 1 "Significant diel vertical shifting appears to occur. " (Jereb & Roper, 2010) (Jereb & Roper, 2010) NA 0 NA 0 1 (McSweeny 1971) 5 "is a predator on Euphausia superba krill" (Jereb & Roper, 2010) "Fish and crustaceans (macroplankton: euphausiids
(incl. Antarctic krill), amphipods, copepods)" (Xavier et al., 2018)		 Jereb & Roper 2010
Xavier et al 2018		 0 0 2 11 "one of the most abundant of Antarctic species of squid and is significant prey for albatrosses, Patagonian toothfish, southern elephant seals and cetaceans" (Jereb & Roper, 2010) Jereb & Roper, 2010
Xavier et al., 2018
Cherel et al. 2017
Imber 1978
Cherel & Duhamel, 2004
Piatkowski & Hagen 1994		 4 0 not gregarious (inferred from photo and video material) NA 0 NA 24 29 96.4 297 NY
Gonatus fabricii Gonatus fabricii 3000 "it is found over a considerable depth range, from surface waters at night to depths of 3000 m during daylight hours (kristensen 1983)" (Hastie et al., 2009). (Jereb & Roper, 2010; Lischka et al., 2019; Hastie et al., 2009; Lischka et al., 2019; Frandsen and Wieland, 2004; Wilborg et al., 1984; Bjorke, 2001; Nesis, 2001; Boyle, 1983; Jereb et al., 2015; Boyle & Rodhouse, 2005) 160 [AL: Younger specimens definitly can be found higher in the water column, but evidence for adult are not clear] "Until 1995, only seven mature specimens of G. fabricii had been recorded and only one speci- men described in detail 􏰁Kristensen, 1981b, 1984; Sennikov et al., 1989). These specimens were recorded as being caught at depths varying from 160 to 2700 m. In 1995, three mature males and two mature females were caught with a pelagic trawl at depths between 270 and 820 m in an area off Andenes, with a bottom depth of 1350 m 􏰁Fig. 1) 􏰁Bjùrke and Hansen, 1996)" (Bjorke, 2001). (Jereb & Roper, 2010; Lischka et al., 2019; Hastie et al., 2009; Lischka et al., 2019; Frandsen and Wieland, 2004; Wilborg et al., 1984; Bjorke, 2001; Nesis, 2001; Boyle, 1983; Jereb et al., 2015; Boyle & Rodhouse, 2005) 1 "Oceanic, Arctic " (Gomes-Pereira et al., 2016) Boyle & Rodhouse, 2005; Gomes-Pereira et al., 2016; Jereb et al., 2015 38 80 "The areas sampled, along with corresponding details of timing and institutions involved, are as follows: (1) Barents
Sea and surroundings from 4◦ E to 77◦ E and as far as 83◦ N (sampled in 2005–2016 by the Polar Research Institute of Marine Fisheries
and Oceanography and Institute of Marine Research); (2) coastal
waters around Greenland, as far as 73◦N on the western side and
67◦N on the eastern side (sampled in 2014–2016 by the Greenland
Institute of Natural Resources); and (3) Canadian side of the Davis
Strait and the Baffin Bay, from 61◦N to 73◦N (sampled in 2016 by
Fisheries and Oceans Canada)" (Golikov et al., 2019).		 (Golikov et al., 2019). 42 80 N to 42 N (Jereb & Roper, 2010) (Jereb & Roper, 2010) 3 1 "This species undertakes vertical migrations and, possibly, horizontal migrations. Kris-
tensen (1977b) reports dispersal of young squid by the West Greenland Current from
zone 1 to zone 2, although no other extensive horizontal migrations are reported. Var-
iation in the barium:calcium ratio in the statoliths of this species suggests that juveniles
inhabit surface waters and that larger specimens move to deeper waters. In addition,
increases in the uranium:calcium and strontium:calcium ratios towards the outer part
of the statolith suggested migration of adult squid into colder water (Zumholz et al.,
2007b). " (Jereb et al., 2015)		 (Jereb et al., 2015) 1 "…the species lives in shoals in the upper- most 80 m of the water column. At increasing size they are found deeper, and as sub-adults and adults they live above the bottom from 200 m downwards, but migrate upwards at night" (Bjorke, 2001). (Bjorke, 2001; Jereb et al., 2015; Frandsen and Wieland, 2004; Golikov et al., 2019; Arkhipkin and Bjorke, 1999; Zumholz and Frandsen, 2006; Boyle, 1983) 1 "…the species lives in shoals in the upper- most 80 m of the water column. At increasing size they are found deeper, and as sub-adults and adults they live above the bottom from 200 m downwards, but migrate upwards at night" (Bjorke, 2001). (Bjorke, 2001; Jereb et al., 2015; Frandsen and Wieland, 2004; Zumholz and Frandsen, 2006; Zumholz et al., 2007; Folkow and Blix, 1999; Boyle, 1983) 1095.8 "As in other squid species (e.g. Illex coindetii, Jereb and Ragonese, 1995; Arkhipkin et al.,
1998), different methods to investigate growth and age give different results for G. fab-
ricii. Analysis of length frequency data suggests a rather long life cycle (2–3 years,
Muus, 1962; Zumholz and Frandsen, 2006)." (Jereb et al., 2015)		 (Lischka et al., 2019; Jereb et al., 2015; Rodhouse and Hatfield, 1990) 365.3 “G. fabricii from Greenland where growth is apparently somewhat slower than in the same species from Norway [largest squid in each population sampled were no more than about 1 year old (Wiborg et al., 1984) , and the maximum age is nearly 2 years (Kristensen, 1983 ). ” (Rodhouse and Hatfield, 1990) (Lischka et al., 2019; Jereb et al., 2015; Rodhouse and Hatfield, 1990) 1095.8 "In Greenland waters, males mature at an age of about 2 years (20-25 cm pen length) whereas females are assumed to mature at an age between 2.5-3 years (Kristensen 1984)" (Frandsen and Wieland, 2004). (Frandsen and Wieland, 2004) 730.5 "In Greenland waters, males mature at an age of about 2 years (20-25 cm pen length) whereas females are assumed to mature at an age between 2.5-3 years (Kristensen 1984)" (Frandsen and Wieland, 2004). (Frandsen and Wieland, 2004) NA 0 NA 0 43 "The stomach contents of gonatus from pelagic trawl hauls were .. studied (WIBORG 1980, 1982). Various organisms, mostly crusta- ceans, were identified. The average frequencies of the most important groups were: Amphipoda 62.8%, Copepoda 33.6%, Chae- tognatha 21.9%, Euphausiacea 7.7%. Larger gonatus had also taken fry of Sebastes sp., Maurolicus muelleri, and small gonatus. The following species were identified: Amphipoda: Parathemisto abyssorum, P. libellula and P. gaudichaudii; Copepoda: Calanus finmarchicus and Pareuchaeta norwegica; Chaetognatha: Sagitta sp. and Eukrohnia sp.; Euphausiacea: Meganyctiphanes norwegica; Pteropoda: Spiratella sp." (Wilborg et al., 1982). Wilborg et al., 1982
Nesis, 1965; Sennikov et al., 1989; Jereb et al., 2015
Bodini et al., 2009
Kristensen, 1984; Jereb et al., 2015		 0 1 Wilborg et al., 1982 6 52 "Gonatus
fabricii is an abundant food source that is exploited by a large number of
marine pred- ators in the Arctic and North Atlantic (see Bjørke 2001 for a
review). These include several species of whale (Globicephala melaena,
Hyperoodon ampullatus, Monodon monoceros, Physeter macro- cephalus, Ziphius
cavirostris) (hjort & Ruud 1929, Grimpe 1925, Nesis 1965, santos et al.
1999, 2001a, 2002), seals (Cysophora cristata, Phoca groenlandica) (potelov et
al. 1997, lydersen et al. 1989), seabirds (Fratercula arctica, Fulmaris
glacialis) (Falk et al. 1992, Garthe et al. 2004), fish (Coryphaenoides
armatus, Gadus morhua, Histiobranchus bathybius, Sebastes marinus, Seriola
dumerili, Reinhardtius hippoglossoides) (Grimpe 1925, Nesis 1965, Matallanas et
al. 1995, Martin & Christiansen 1997, Dawe & Brodziak 1998) and other
squid (Illex illecebrosus) (Amaratunga 1983) (Table 6). in certain offshore
areas of the North Atlantic, particularly around the Faroes and Norwegian sea,
puffins (Fratercula arctica) have been observed feeding almost exclusively on
G. fabricii (Falk et al. 1992)" (Hastie et al., 2009 )		 (Hovde et al., 2002) (Garthe
et al., 2004) (Haug
et al., 2007) (Hastie et al., 2009 ) (Wilborg et al., 1982) (Finley
and Gibb, 1982) (Lick
and Piatkowski, 1998) (Enoksen
et al., 2017) (Bjorke,
2001) (Haug
et al., 2004) (Michalsen
and Nedreaas, 1998) (Xavier
et al., 2018) (Jereb
et al., 2015) 		5 3 a "shoaling squid" (Kristensen in Boyle 1983:171) Bjorke, 2001; Kristensen in Boyle 1983:171 1 a "shoaling squid" (Kristensen in Boyle 1983:171) Bjorke, 2001; Kristensen in Boyle 1983:171 2 0 2 1 "Current knowledge suggests that G. fabricii reproduces in only
a few specific geographic areas (e.g. the northeastern part of the
Greenland Sea, the border between the Greenland and Norwegian
Seas, the northeastern and southernmost parts of the Norwegian
Sea and the southwestern part of the Davis Strait), but this needs to
be clarified (Bjørke, 1995, 2001; Bjørke & Gjøsaeter, 1998). Some
authors have indicated that this species may reproduce throughout
its distributional range (Nesis, 1965, 1971, 1987)" (Golikov et al., 2019).		 (Golikov et al., 2019). 1 "maturing and spawning females undergo rapid degradation of tissue of the mantle and arms into a gelatinous consistency; tentacles and arm suckers are lost, and normal, active locomotion is reduced. Egg masses are held in the arms while the embryos develop. After eggs hatch into paralarvae, the female dies." (Jereb & Roper, 2010) (Jereb & Roper, 2010) 0 "maturing and spawning females undergo rapid degradation of tissue of the mantle and arms into a gelatinous consistency; tentacles and arm suckers are lost, and normal, active locomotion is reduced. Egg masses are held in the arms while the embryos develop. After eggs hatch into paralarvae, the female dies." (Jereb & Roper, 2010) (Jereb & Roper, 2010) 87 56 101.4 110 NY
Grimalditeuthis bonplandi Grimalditeuthis bonplandi 2046 “BS 2010–2046 m” (Urbano and Hendrickx, 2018) (Judkins & Vecchione, 2020; Urbano and Hendrickx, 2018) 600 "Fifty-three Grimalditeuthis bonplandi (6–84 mm ML) were analyzed. They were
found from the surface to 1500 m during day and night
with the majority of individuals found between 600 and
1500 m. We found no evidence of vertical migration but
an ontogenic shift for this species is evident, with larger
individuals found in the lower meso- and upper bathypelagic
zones " (Judkins & Vecchione, 2020)		 (Judkins & Vecchione, 2020; Urbano and Hendrickx, 2018) 1 "Mesopelagic to bathypelagic." (Jereb & Roper, 2010) Chesalin & Zuyev, 2002;
Jereb & Roper, 2010		 85 45 45 N to 40 S (Jereb & Roper, 2010) (Jereb & Roper, 2010) -40 45 N to 40 S (Jereb & Roper, 2010) (Jereb & Roper, 2010) 4 1 "an ontogenic shift for this species is evident, with larger individuals found in the lower meso- and upper bathypelagic zones" (Judkins & Vecchione, 2020) (Judkins & Vecchione, 2020) 1 1 1 "These tentacle club movements superficially resemble the movements of small marine organisms and suggest the possibility that G. bonplandi uses aggressive mimicry by the tentacle clubs to lure prey, which we find to consist of crustaceans and cephalopods. In the darkness of the meso- and bathypelagic zones the flapping and undulatory movements of the tentacle may: (i) stimulate bioluminescence in the surrounding water, (ii) create low-frequency vibrations and/or (iii) produce a hydrodynamic wake. Potential prey of G. bonplandi may be attracted to one or more of these as signals. This singular use of the tentacle adds to the diverse foraging and feeding strategies known in deep-sea cephalopods" (Hoving et al 2013) Hoving et al 2013 1 1 1 inking (Bush & Robison, 2007, Table 1) Bush & Robison, 2007 4 "Cephalopods, crustaceans, Hoving et al. (2013b)" (Elsevier, 2014) Hoving et al 2013
Rodhouse and Piatkowski (1995)		 0 0 3 5 Found
in the stomach of the blue shark (Prionace glauca) (Markaida &
Sosa-Nishizaki, 2010)		 (Markaida & Sosa-Nishizaki, 2010) (Urbano & Hendrickx, 2018) (Tsuchiya and Sawadaishi 1997) 3 0 not gregarious (inferred from photo and video material) NA 0 NA 5 5 27.8 67 NY
Haliphron atlanticus Haliphron atlanticus 6787 "Reported to occupy a depth range in open-ocean from the surface to at least 1 260 m, over depths of up to 6787 m. Collected in bottom trawls on continental shelves and slopes at depths of 100 to 3173 m" (Jereb et al., 2014) (Judkins & Vecchione, 2020; Jereb et al., 2014; Vecchione & Roper, 1991, p. 438; Goncalves, 1991; Jereb et al., 2014) 0 "Twenty-nine Haliphron atlanticus were collected from the surface
layer down to 1200 m. Mantle lengths ranged from 4 to 32 mm
with no evidence of diel vertical migration. Larger H. atlanticus
were found in the upper layers " (Judkins & Vecchione, 2020)		 (Judkins & Vecchione, 2020; Jereb et al., 2014; Vecchione & Roper, 1991, p. 438; Goncalves, 1991; Jereb et al., 2014) 2 **borderline** JM: gelatinous; needs to float so likely pelagic. "Collected in bottom trawls on continental shelves and slopes at depths of 100 to 3 173 m. It has been proposed that Haliphron atlanticus may not be entirely pelagic, but might pass only relatively short periods of its life cycle in the open waters, soon returning to a life at the bottom especially on continental slopes" (Jereb et al., 2014); "Haliphron atlanticus has a wide geographical distribution, and while it occurs in mesopelagic and bathypelagic waters of the open ocean, it is often associated with continental slopes, both in the pelagic water column as well as close to the bottom16." (Hoving & Haddock 2017) Jereb et al., 2014; Hoving & Haddock 2017 128 83 "North Atlantic, but there are two records from the Norwegian coast at 66.3°N–68.3°N" (Xavier et al., 2018) (Xavier et al., 2018) -45 68 N to 45 S (Jereb et al., 2014) (Jereb et al., 2014) 3 1 "The overall size distribution showed evidence of progressive growth throughout the course of the year, with peak body sizes occurring in the fall of 2012 (Fig. 5b) and decreasing slightly in 2014. Brooding has been reported in H. atlanticus (Young 1995) and juveniles are thought to live in near-surface waters, descending into deeper water as they grow and mature (Nesis 1987; Young 1995)" (Shea et al. 2017) (Shea et al. 2017; Judkins & Vecchione, 2020) 2 2 1 holding and possibly preying on jellyfish Periphyllopsis braueri "caught within the oral web of the octopus." (Hunt et al. 2019) Hunt et al. 2019 1 "this octopod is capable of capturing prey since the crop of our specimen was filled with shrimps whose heads had been neatly severed" (Nixon & Young 2003) Nixon & Young 2003 2 1 1 male H. atlanticus holding a jellyfish Pelagia noctiluca, Terceira Island, Azores “Throughout the whole record of the encounter (2:26 mins), the male octopod retained its hold on the jelly, in an oral-to-oral surface orientation…The octopod actively maintained its position up in the water column, a couple of meters from the bottom and clear of any structure, by jetting water from its funnel. Little to no activity was observed from the jelly..The orientation of the interaction and the considerably larger size (and presumed mass) of the octopod seems inconsistent with camouflage, shelter, and/or transportation of the octopod by the jelly…The interaction between the octopod and divers, namely by rotating the jelly towards what we presume to be a perceived threat, supports a potential defensive use of the hijacked jelly. Thus, we argue that the octopod seems to be using the jelly to protect itself…Alternatively, if this footage solely captured the initial stages of a ‘Burglary’ interaction…Such ‘Burglary’ behavior is supported primarily by the otherwise surprising orientation of the animals: most of the jelly’s tentacles were available to the octopod and fragments could, theoretically, be aligned with its arms (as in Tremoctopus) with relative ease. The orientation also likely renders the jelly incapable of escaping until the octopod releases it…we argue that it is unlikely that the footage recorded refers exclusively to a predatory event because of the unwillingness of the octopod to release the jelly.” (Rosa et al. 2019) Rosa et al. 2019 1 (O'Shea, 2004; Jereb et al., 2014) 5 "atlanticus is reported to feed on crustaceans and cephalopods. " (Jereb et al., 2014) Jereb et al 2015
Xavier et al 2013
Hoving & Haddock 2017
Willassen (1986) and O’Shea (2004b)		 0 0 3 10 "It
has been reported in the stomach contents of sperm whales in the northeastern
Atlantic North Sea region [32], the eastern Atlantic Canary Islands [34],
southern Australia [33], and Patagonia of South America [31]…Haliphron
atlanticus is also typically present in small to moderate numbers in the
stomach contents of blue sharks, such as near the Azores in the North Atlantic
[35], of California [38] and Brazil [37], and in the Kuroshio Extension
[36]…their foraging depths can directly overlap with the depth where the H.
atlanticus was seen in the present study and where smaller individuals live
[13]….has been detected in the diets of the Hawaiian monk seal, Monachus
schauinslandi [81], and South Shetland Island Weddell seals, Leptonychotes
weddellii" (Miller et al. 2018)		 (Fernandez et al., 2014) (Markaida & Sosa-Nishizaki, 2010) (Laptikhovsky et al. 2020) (Chua et al. 2019) (Cherel et al., 2009) (Konan et al. 2018) (Miller et al. 2018) (Imber, 1996) (Clarke, 1986) 4 0 not gregarious (inferred from photo and video material) 2 1 1 "The interaction between the octopod and divers, namely by rotating the jelly towards what we presume to be a perceived threat, supports a potential defensive use of the hijacked jelly. Thus, we argue that the octopod seems to be using the jelly to protect itself." (Rosa et al. 2019). "The manner in which Haliphron holds the medusa in our observations is very similar to the description by Heeger et al.31. In Haliphron the medusa was held by the external bell with the oral part open within the folds of the arms, and the octopus’s beak sometimes protruding into the subumbrellar space. Therefore, in addition to feeding directly on jellyfishes, Haliphron may target the stomach contents of the medusa, or even use the medusa as a tool to obtain more nutritious prey that are captured by the fringe of tentacles clasped within the octopus arms" (Hoving & Haddock 2017) Rosa et al. 2019; Hoving & Haddock 2017 1 1 "eggs are incubated in large clusters in the crown of the female’s arms." (Laptikhovsky & Salman, 2002) (Laptikhovsky & Salman, 2002) 0 "eggs are incubated in large clusters in the crown of the female’s arms." (Laptikhovsky & Salman, 2002) (Laptikhovsky & Salman, 2002) 38 27 160.2 NY
Hapalochlaena fasciata Hapalochlaena fasciata 20 "Depth range from 0 to at least 20 m" (Jereb et al., 2014) (Jereb et al., 2014) 0 "Depth range from 0 to at least 20 m" (Jereb et al., 2014) (Jereb et al., 2014) 2 "Occurs on intertidal and shallow rocky reefs." (Jereb et al., 2014) Jereb et al., 2014;
Norman and Reid 2000:56		 13 -24 24 S to 37 S (Jereb et al., 2014) (Jereb et al., 2014) -37 24 S to 37 S (Jereb et al., 2014) (Jereb et al., 2014) 1 121.8 aquarium "The hatchlings were reared in the laboratory for 4 months, at the end of which time they were sexually mature (35-40 mm ML)." (Nixon & Young 2003) (Nixon & Young 2003) 121.8 aquarium "The hatchlings were reared in the laboratory for 4 months, at the end of which time they were sexually mature (35-40 mm ML)." (Nixon & Young 2003) (Nixon & Young 2003) 1 1 1 "It feeds on crustaceans and fish, using a strong toxin to paralyse large prey." (Norman and Reid 2000 – Book, p56) Norman and Reid 2000 – Book, p56
Nixon & Young 2003		 4 3 1 "One of the species of cephalopods sheltering within these seagrass beds is the venomous blue-lined octopus, Hapalochlaena fasciata (Norman and Reid 2000). This visually cryptic animal hides among tide pools, blending in with its surroundings via pigmented chromatophore organs. When threatened, the animal flashes bright blue rings and lines, which act as a warning to potential predators (Tranter and Augustin 1973; Mathger et al. 2008)." (Townsend et al. 2012) Townsend et al. 2012 1 inferred from H. maculosa (Tranter & Augustine 1973) and lunulata (Mathger et al. 2012) Tranter & Augustine 1973; Mathger et al. 2012 1 “An additional defensive role is suggested by the occurrence of TTX in ink. This novel discovery in three of seven H. lunulata and the high concentration in at least one individual are notable…Ink sacs in the genus HapalochlaeNAare highly reduced (Robson, 1929) and were long-thought to have been usurped by the powerful chemical defense of TTX (Roper and Hochberg, 1988; Stranks and Lu, 1991; Tranter and Augustine, 1973). Still, inking behavior has been observed in some adult blue-ringed octopuses (H. lunulata, H. sp. 3, and H. sp. 5; Huffard and Caldwell, 2002; Norman, 2000). While H. lunulata ink was not effective as a defense in two incidents with one mantis shrimp predator, Odontodactylus scyllarus (Huffard and Caldwell, 2002), variation in individual TTX concentrations and potential resistance of the predator species may have affected the outcome in these two cases. The high concentration of TTX in the ink of H. lunulata #9 and the presence of TTX in the ink of H. lunulata #6, which lacked detectable TTX in the PSG, suggest that occurrence there is not accidental.” (Williams & Caldwell 2009) Williams & Caldwell 2009 2 "“Hatchling HapalochlaeNAare able to expel ink: H. fasciata (Hoyle, 1886) (Tranter and Augustine 1973 [taxonomy according to Stranks and Lu 1991])…However, H. fasciata does not ink after they are 4 weeks old (Tranter and Augustine 1973)…Tranter
and Augustine (1973) also reported that H.
fasciata does not eject ink as adults” (Huffard & Caldwell 2002)"		 (Hoyle, 1886; Stranks and Lu, 1991; Huffard & Caldwell 2002; Jereb et al., 2014) 1 "This is the first evidence of the antipredator function of TTX. While omnivorous green turtle hatchlings have been observed attempting to feed on small, non-venomous octopuses, Octopus bocki (Caldwell 2005), we propose that these are two cases of healthy, adult herbivorous green sea turtles (C. mydas) accidentally consuming cryptic blue-lined octopuses (H. fasciata) while feeding on the seagrass meadows in which the octopus shelters. These accidental ingestions resulted in lethal TTX envenomations by the blue-lined octopuses, delivered actively through biting the esophageal tissue. The TTX subsequently immobilized the turtle, preventing it from raising its head above the water to breathe, filling its lungs with water and resulting in a toxin-induced drowning." (Townsend et al. 2012); “The results of this study highlight several characteristics regarding the defensive use of TTX by H. cf. fasciata….Considering that H. cf. fasciata and moray eel were separated by a clear acrylic plate with small holes, this species actively secretes TTX from its body surface when it recognizes the presence of a predator by visual or olfactory cues. When the moray eel approached the octopus in this experiment, the characteristic blue ring and black stripes that develop during vigilance (Mäthger et al. 2012) indicate that the octopus recognized the predator as a danger.” (Yamate et al. 2024) Townsend et al. 2012; Yamate et al. 2024 2 predates White's seahorse Hippocampus whitei in Port Stephens marine estuary off Australia (Manning et al. 2018) Jereb et al 2014
Manning et al. 2018		 0 0 2 0 not gregarious (inferred from photo and video material) 1 0 1 1 "The female Blue-lined octopus plaits her large
eggs into strings and carries them in her web until they
hatch." (Norman and Reid 2000 – Book, p56)		 (Norman and Reid 2000 – Book, p56) 0 "The female Blue-lined octopus plaits her large
eggs into strings and carries them in her web until they
hatch." (Norman and Reid 2000 – Book, p56)		 (Norman and Reid 2000 – Book, p56) 10 11 54.9 49.7 36 28 C22
Helicocranchia papillata Helicocranchia pfefferi 2000 "Vertical distribution of paralarvae and juveniles occurs in the upper 300 m, followed by ontogenetic descent of individuals into the mesopelagic zone with growth. Adults mature in the bathypelagic waters of 2 000 m or more; shallow depth captures of paralarvae and nearly spent females in the upper 100 m point to an epipelagic spawning habit." (Jereb & Roper, 2010) (Jereb & Roper, 2010) 2000 [AL: We focus on non post-spawning moribund adults] "Vertical distribution of paralarvae and juveniles occurs in the upper 300 m, followed by ontogenetic descent of individuals into the mesopelagic zone with growth. Adults mature in the bathypelagic waters of 2 000 m or more; shallow depth captures of paralarvae and nearly spent females in the upper 100 m point to an epipelagic spawning habit." (Jereb & Roper, 2010) (Jereb & Roper, 2010) 1 "Vertical distribution of paralarvae and juveniles occurs in the upper 300 m, followed by ontogenetic descent of individuals into the mesopelagic zone with growth. Adults mature in the bathypelagic waters of 2 000 m or more; shallow depth captures of paralarvae and nearly spent females in the upper 100 m point to an epipelagic spawning habit." (Jereb & Roper, 2010) Jereb & Roper, 2010 25 35 " The geographical distribution of H. papillata occupies a broad portion of the western, central and
eastern subtropical North Atlantic Ocean; Caribbean Sea and Gulf of Mexico, northeastern African waters." (Jereb & Roper, 2010)		 (Jereb & Roper, 2010) 10 " The geographical distribution of H. papillata occupies a broad portion of the western, central and
eastern subtropical North Atlantic Ocean; Caribbean Sea and Gulf of Mexico, northeastern African waters." (Jereb & Roper, 2010)		 (Jereb & Roper, 2010) 3 1 "Vertical distribution of paralarvae and juveniles occurs in the upper 300 m, followed by ontogenetic descent of individuals into the mesopelagic zone with growth. Adults mature in the bathypelagic waters of 2 000 m or more; shallow depth captures of paralarvae and nearly spent females in the upper 100 m point to an epipelagic spawning habit." (Jereb & Roper, 2010) (Jereb & Roper, 2010) NA 0 NA 0 0 0 0 not gregarious (inferred from photo and video material) NA 0 NA 1 1 7.2 NY
Heteroteuthis dispar Heteroteuthis dispar 800 [AL: Caugth at lower depth but no clear evidence of adult caught deeper] In hauls from 50-800 they only inhabited 600-800 m (Quetglas et al., 2000) (Salman et al., 2003; Jereb & Roper, 2005; Quetglas et al., 2000; Sifner et al., 2005; Rotermund & Guerrero-Kommritz 2010; Lefkaditou et al. 1999; Villanueva 1992; Roper & Young, 1975) 100 Caught between 100 m and 650 m (Salman et al., 2003) (Salman et al., 2003; Jereb & Roper, 2005; Quetglas et al., 2000; Sifner et al., 2005; Rotermund & Guerrero-Kommritz 2010; Lefkaditou et al. 1999; Villanueva 1992; Roper & Young, 1975) 2 **edge case** JM: pelagic lifestyle. “Small cephalopods of the genus Heteroteuthis are the most pelagic members in the family Sepiolidae…The species seems to be widely distributed throughout the watercolumn. In two studies performed in the Aegean Sea, H. dispar was found to be the most common cephalopod in pelagic samplings up to 650 m of depth (Salman et al. 2003; Lefkaditou et al. 1999). Nesis (1993) found members of the genus Heteroteuthis, although broadly distributed in the open ocean, more common near seamounts and submarine ridges. Roper (1974) reported H. dispar migrating to the surface at night. In summarising the life history of H. dispar, Nesis (1985) stated that eggs are laid on the bottom of the slope (500–1,000 m), and hatchlings ascend to epipelagic layers. Immature H. dispar occur at 25–300 m in the night and at 150–500 m during the day. Maturing animals gradually descend to deeper waters, and mature H. dispar are distributed in near-bottom layers. Spawning occurs on the seabed.” (Hoving et al. 2008) Hoving et al. 2008 51 52 52 N to 1 N (Jereb & Roper, 2005) (Jereb & Roper, 2005) 1 52 N to 1 N (Jereb & Roper, 2005) (Jereb & Roper, 2005) 2 730.5 2 yrs (Rotermund & Guerrero-Kommritz 2010) (Rotermund & Guerrero-Kommritz 2010; Nixon & Young, 2003:81) 730.5 2 yrs (Rotermund & Guerrero-Kommritz 2010) (Rotermund & Guerrero-Kommritz 2010; Nixon & Young, 2003:81) 365.3 maturing "during the second year" (Nixon & Young, 2003:81) (Nixon & Young, 2003:81) 365.3 maturing "during the second year" (Nixon & Young, 2003:81) (Nixon & Young, 2003:81) NA 0 2 2 1 "the production of light by midwater squids is believed to be a means to conceal themselves from predators by counter-illumination (Young & Roper 1976; Young et al. 1980)." (Bello 1997) Bello 1997
Nixon & Young 2003		 1 "among cephalopods, only species of the sepiolid genus Heteroteuthis are known to release luminous Xuid with their ink (Herring 1977; Dilly and Herring 1978; P. Herring personal communication)." (Bush & Robison, 2007) Bush & Robison, 2007
Hanlon & Messenger, 2018, p. 130		 1 "Ink is sometimes discharged at the same time as the luminous secretion into the sea. We have no evidence of whether the ink can be discharged without the luminous secretion…The luminous gland of Heteroteuthis does not contain bacteria. It is undoubtedly the source of the luminous secretion produced by the squid, and an alternative mode of production of the light must be found. Our observations suggest that the luminous material is secreted by the cells that surround the lumen of the gland." (Dilly & Herring 1978) (Dilly & Herring 1978; Jereb & Roper, 2005) 0 0 10 "It
is a dominant presence in the stomachs of albacore (Thunnus alalunga) taken
from the Adriatic Sea (Bello 1999) and of giant red shrimp (Aristaeomorpha
foliacea) from the Strait of Sicily (Bello and Pipitone 2002). H. dispar plays
also an important role in the diet of benthic bathyal elasmobranchs, such as
Galeus melastomus and Etmopterus spinax (Sartor and De Ranieri 1995).
Additionally, H. dispar has been found in stomachs of the swordfish (Xiphias
gladius) (Bello 1991; Villanueva 1995), Risso’s dolphin (Grampus griseus)
(Würtz et al. 1992), and the deep-sea shark Centroscymnus coelolepis
(Villanueva 1992)." (Hoving
et al. 2008)		 (Blanco et al., 2006) (Jereb & Roper, 2005) (Kousteni et
al. 2018) (Battaglia et al. 2013) (Romeo et al. 2012) (Hoving et al. 2008) (Bello
1997) (Bello 1990)		 5 3 "Adults live frequently in groups in the lower epipelagic and mesopelagic zones, most commonly in depths between 200 and 300 m" Jereb & Roper, 2005: 199 1 "Adults live frequently in groups in the lower epipelagic and mesopelagic zones, most commonly in depths between 200 and 300 m" Jereb & Roper, 2005: 199 NA 0 "In living animals, tips of the arms and fins are translucent except for some brownish chromatophores placed distally on the fins (Bello & Biagi, 1995). Animals are able to illuminate their ventral side in a blue-green glow. The maximum wave-length of the photophore lies between 485 and 490 nm (Nixon et al., 2003)." (Rotermund & Guerrero-Kommritz 2010) NA "Sucker scars, present on the head and mantle in both females and males match the enlarged suckers on the third arms of the male, which indicates that these arms are used to hold the female during mating (Orsi-Relini 1995). The presence of scars on males may be due to agonistic behaviour between competing males, or males may capture any specimens within their reach regardless their sex and only start mating when the captured specimen is a female" (Hoving et al. 2008) 36 26 42.642 29.394 12.6 20 16 11 W-W-NY
Histioteuthis miranda Histioteuthis miranda 1200 "Capture data suggest that this is the most abundant histioteuthid species encountered throughout most of its normal range. It occurs abundantly at 700 to 900 m off South Africa, where a large breeding population is located. It ranges from subsurface waters to at least 1 200 m; ontogenetic descent exists as larger subadults and adults generally are captured deeper than juveniles, 700 to 1 200 m on or close to the bottom; elsewhere subadults and mature males and females were taken both day and night at or near the bottom at 400 to 1 200 m. Many specimens of H. miranda have been caught frequently in deep-water, bottom-fishing, lobster trawls at 300 to 600 m off the North West Shelf of Australia and the Great Barrier Reef, suggesting an association with the sea floor at some time during their life cycle." (Jereb & Roper, 2010) (Jereb & Roper, 2010; Hoving et al 2010) 300 "Capture data suggest that this is the most abundant histioteuthid species encountered throughout most of its normal range. It occurs abundantly at 700 to 900 m off South Africa, where a large breeding population is located. It ranges from subsurface waters to at least 1 200 m; ontogenetic descent exists as larger subadults and adults generally are captured deeper than juveniles, 700 to 1 200 m on or close to the bottom; elsewhere subadults and mature males and females were taken both day and night at or near the bottom at 400 to 1 200 m. Many specimens of H. miranda have been caught frequently in deep-water, bottom-fishing, lobster trawls at 300 to 600 m off the North West Shelf of Australia and the Great Barrier Reef, suggesting an association with the sea floor at some time during their life cycle." (Jereb & Roper, 2010) (Jereb & Roper, 2010; Hoving et al 2010) 2 **edge case** JM: pelagic lifestyle. “Small cephalopods of the genus Heteroteuthis are the most pelagic members in the family Sepiolidae…The species seems to be widely distributed throughout the watercolumn. In two studies performed in the Aegean Sea, H. dispar was found to be the most common cephalopod in pelagic samplings up to 650 m of depth (Salman et al. 2003; Lefkaditou et al. 1999). Nesis (1993) found members of the genus Heteroteuthis, although broadly distributed in the open ocean, more common near seamounts and submarine ridges. Roper (1974) reported H. dispar migrating to the surface at night. In summarising the life history of H. dispar, Nesis (1985) stated that eggs are laid on the bottom of the slope (500–1,000 m), and hatchlings ascend to epipelagic layers. Immature H. dispar occur at 25–300 m in the night and at 150–500 m during the day. Maturing animals gradually descend to deeper waters, and mature H. dispar are distributed in near-bottom layers. Spawning occurs on the seabed.” (Hoving et al. 2008) Hoving & Lipiński 2009; Jereb & Roper, 2010; Roeleveld et al. 1992; Hoving et al. 2010 52 12 "This study finds evidence of their presence in the eastern Arabian Sea close to the Lakshadweep archipelago and
associated submarine mounts (Perumal Par, Suheli Par,
Cherbaniani Reef and Byragmore Reef). Voss et al. (1998)
first reported H. miranda from the western Arabian Sea
(~12° N, 52° E), which is about 1240 nm northwest of the
present record" (Sajikumar et al 2018)		 (Sajikumar et al 2018) -40 10 S to 40 S (Jereb & Roper, 2010) (Jereb & Roper, 2010) 4 1 "Capture data suggest that this is the most abundant histioteuthid species encountered throughout most of its normal range. It occurs abundantly at 700 to 900 m off South Africa, where a large breeding population is located. It ranges from subsurface waters to at least 1 200 m; ontogenetic descent exists as larger subadults and adults generally are captured deeper than juveniles, 700 to 1 200 m on or close to the bottom; elsewhere subadults and mature males and females were taken both day and night at or near the bottom at 400 to 1 200 m. Many specimens of H. miranda have been caught frequently in deep-water, bottom-fishing, lobster trawls at 300 to 600 m off the North West Shelf of Australia and the Great Barrier Reef, suggesting an association with the sea floor at some time during their life cycle." (Jereb & Roper, 2010) (Jereb & Roper, 2010) 730.5 "…while Histioteuthis miranda Berry, 1918, from the same area may live for up to 2 years (Hoving and Lipinski, 2009)" (Elsevier, 2014). (Elsevier, 2014) 730.5 "…while Histioteuthis miranda Berry, 1918, from the same area may live for up to 2 years (Hoving and Lipinski, 2009)" (Elsevier, 2014). (Elsevier, 2014) NA 0 NA 0 1 "Crustaceans, Clarke (1980)" (Elsevier, 2014). Clarke (1980) 0 0 1 6 both
juveniles and adults made up 1.9% of cephalopod speciesfound in stomach of
juvenile male Pygmy Sperm Whale stranded in SE Tasmania (Beasley et al. 2013)		 "Jereb & Roper, 2010"
"Beasley et al. 2013"
"Wang et al. 2002"
"Sekiguchi et al. 1996"
"Clarke et al 1993"
"Imber, 1996"		 3 0 not gregarious (inferred from photo and video material) NA 0 NA 6 7 131.6 60 NY
Illex coindetii Illex coindetii 1100 "Occurs from the surface to about 1100 m" (Lauria et al., 2016) Lauria et al., 2016
Vafidis et al 2008		 0 "In the Mediterranean Sea I. coindetii has been recorded from surface waters to over 700 m (i.e., 776 m, South Aegean Sea; Lefkaditou et al., 2003), with highest densities found between 100–200 and 400–600 m (e.g., Tursi and D’Onghia, 1992; Jereb and Ragonese, 1995; Salman et al., 1997)" (Arkhipkin et al., 2015) Arkhipkin et al., 2015 2 **edge case** JM Feb 28: definitely pelagic. "is pelagic and semi-demersal" (Salman et al., 2003). "Illex coindetii is the most common benthic ommastrephid squid exploited by bottom trawl in the Mediterranean Sea" (Lefkaditou et al. 2008) Lefkaditou et al. 2008; Salman et al., 2003 80 60 Western Atlantic 5 N to 40 N and eastern Atlantic 20 S to 60 N (Arkhipkin et al., 2015) (Arkhipkin et al., 2015) -20 Western Atlantic 5 N to 40 N and eastern Atlantic 20 S to 60 N (Arkhipkin et al., 2015) (Arkhipkin et al., 2015) 2 1 “It is suggested that Illex populations do not have any specific areas of aggregation during summer months. During fall and winter months, species growth rate increases and as a highly mobile and opportunistic species, they migrate offshore to take advantage of upwelling regions and associated plankton blooms (Boyle, 1983). Winter offshore upwelling events in the study area occur at locations around Antikithira Strait and southwest of Crete Island (Valavanis et al., 1999), mainly due to seasonal strong winds and associated gyres in the region (Theocharis et al., 1993). During spring months with spring spawning season approaching, species start their spawning migration in a southward direction (Amaratunga, 1981; Dawe et al., 1981; Rathjen, 1981) to find warmer spawning and egg development temperature ranges (Boletzky et al., 1973).” (Valavanis et al. 2002) but see "The lack of evidence of seasonal migrations is in agreement with the available
preliminary information on I. coindetii of the Sicilian Channel (Jereb & Ragonese, 1991),
and could be a result of the characteristics of this area of the Mediterranean." (Jereb & Ragonese, 1995) [N=11700]		 (Valavanis et al. 2002) (Jereb & Roper, 2010) 1 "Juveniles and adults share the same depth ranges in some Mediterranean areas (Sanchez et al., 1998; Ceriola et al., 2006), though juveniles/small specimens show a major concentration in waters shallower than 200 m. Adults undergo vertical migrations from the bottom to the upper layers at night and seasonal migrations have been observed in the western and central Mediterranean waters (Mangold-Wirz, 1963; Soro and Paolini, 1994; Sanchez et al., 1998; Gentiloni et al., 2001) with the bulk of the population approaching shallow waters (70–150 m) in spring, to spread again over a wider bathymetric range in autumn." (Arkhipkin et al., 2015) (Arkhipkin et al., 2015) (Salman, 2017) 1 "The species appears to be associated with the bottom during the day, dispersing into mid-depths at night" (Cairns, 1976) (Cairns, 1976) Arkhipkin et al., 2015) 547.875 "Based on length frequency analyses, the maximum lifespan of I. coindetii from different geographic areas has been estimated to be 12–18 months, whereas direct age determi-
nation by statolith reading has indicated lifespans as short as 6 months (Table 15.2). Length frequency distributions for cephalopod species of interest to fisheries are gen-
erally polymodal, but it is difficult to identify microcohorts, and growth estimates by
means of length frequency methods are difficult to make (Sánchez, 1984; Caddy, 1991;
Jereb and Ragonese, 1995; Arvanitidis et al., 2002). Therefore, direct age determination
methods are applied more frequently. Despite the acknowledged validity of the meth-
odology (Jereb et al., 1991; Jackson, 1994; Ceriola and Milone, 2007), several authors
have advised caution in interpreting age values from statolith readings (e.g. Lipiński
and Durholtz, 1994; González et al., 2000; Bettencourt and Guerra, 2001). " (Jereb et al., 2015)		 (Jereb et al., 2015) 182.625 "Based on length frequency analyses, the maximum lifespan of I. coindetii from different geographic areas has been estimated to be 12–18 months, whereas direct age determi-
nation by statolith reading has indicated lifespans as short as 6 months (Table 15.2). Length frequency distributions for cephalopod species of interest to fisheries are gen-
erally polymodal, but it is difficult to identify microcohorts, and growth estimates by
means of length frequency methods are difficult to make (Sánchez, 1984; Caddy, 1991;
Jereb and Ragonese, 1995; Arvanitidis et al., 2002). Therefore, direct age determination
methods are applied more frequently. Despite the acknowledged validity of the meth-
odology (Jereb et al., 1991; Jackson, 1994; Ceriola and Milone, 2007), several authors
have advised caution in interpreting age values from statolith readings (e.g. Lipiński
and Durholtz, 1994; González et al., 2000; Bettencourt and Guerra, 2001). " (Jereb et al., 2015)		 (Jereb et al., 2015) 285 "Age at maturation varies between 120 and 271 d in males and between 120 and 285 d
in females, depending on the geographic area and season considered (González et al.,
1996a; Arkhipkin et al., 1998). Individuals of this species mature at a wide range of
sizes. Although size at maturity shows some degree of geographic variation in both
sexes (e.g. Arvanitidis et al., 2002; Hernández-García, 2002a), males mature at a lower
minimum size than females." (Jereb et al., 2015)		 (Jereb et al., 2015) 120 "Age at maturation varies between 120 and 271 d in males and between 120 and 285 d
in females, depending on the geographic area and season considered (González et al.,
1996a; Arkhipkin et al., 1998). Individuals of this species mature at a wide range of
sizes. Although size at maturity shows some degree of geographic variation in both
sexes (e.g. Arvanitidis et al., 2002; Hernández-García, 2002a), males mature at a lower
minimum size than females." (Jereb et al., 2015)		 (Jereb et al., 2015) NA 0 NA 0 20 "Main prey are fish and crustaceans." (Lauria et al., 2016) Jereb & Roper, 2010
Luaria et al 2016
Castro & Hernandez-Garcia, 1995		 0 1 Jereb & Roper, 2010 4 28 (Jereb et al., 2015) "Table 15.6. Known predators of Illex coindetii in the Northeast Atlantic and Mediterranean Sea." Taxon Species References
Cephalopoda: European squid (Loligo vulgaris) Dawe and Brodziak (1998)
European flying squid (Todarodes sagittatus) Dawe and Brodziak (1998)
Chondrichthyes: Black-mouthed dogfish (Galeus melastomus) Valls et al. (2011)
Lesser spotted dogfish (Scyliorhinus canicula) Kabasakal (2002)
Sharpnose sevengill shark (Heptranchias perlo) Henderson and Williams (2001)
Shortfin mako (Isurus oxyrinchus) Maia et al. (2006)
Smooth-hound (Mustelus mustelus) Kabasakal (2002)
Thornback ray (Raja clavata) Kabasakal (2002), Farias et al. (2006), Šantić et al. (2012)
Osteichthyes Albacore (Thunnus alalunga) Consoli et al. (2008), Romeo et al. (2012)
Atlantic bluefin tuNA(Thunnus thynnus) Karakulak et al. (2009), Romeo et al. (2012), Battaglia et al. (2013)
Blonde ray (Raja brachyura) Farias et al. (2006)
Blue whiting (Micromesistius poutassou) Macpherson (1978)
Common dolphinfish (CoryphaeNAhippurus) Massutí et al. (1998)
Conger eel (Conger conger) Lordan et al. (1998b)
Greater forkbeard (Phycis blennoides) Morte et al. (2002)
Mediterranean spearfish (Tetrapturus belone) Castriota et al. (2008), Romeo et al. (2009, 2012)
Saithe (Pollachius virens) Lordan et al. (1998b)
Smooth lanternshark (Etmopterus pusillus) Xavier et al. (2012)
Swordfish (Xiphias gladius) Bello (1985), Moreira (1990), Salman (2004), Peristeraki et al. (2005), Romeo et al. (2009, 2012)
Yellowfin tuNA(Thunnus albacares) Dragovich (1970)
Cetacea Bottlenose dolphin (Tursiops truncatus) González et al. (1994a), Santos et al.
(1997)
Common dolphin (Delphinus delphis) González et al. (1994a), Silva (1999a)
Long-finned pilot whale (Globiocephala melas) González et al. (1994a)
Risso’s dolphin (Grampus griseus) Carlini et al. (1992), González et al. (1994a), Santos et al. (1997), Blanco et al. (2006), Bearzi et al. (2011)
Striped dolphin (Stenella coeruleoalba) Würtz and Marrale (1993), Alessandri et al. (2001) 		(Blanco et al., 2006) (Arkhipkin et al., 2015) (Jereb &
Roper, 2010) (Dawe & Brodziak 1998) Quetglas et al. (1998) (Kousteni et al.
2018) (Battaglia et al. 2013) (Consoli et al. 2008) (Bello 1997) (Velasco et
al., 2001) (Pierce et al., 2010)) (Romeo et al. 2009) (Jereb et al., 2015) (Varela
et al 2018) (Abid et al 2018) (Battaglia et al 2013) (Castriota et al 2008) (Henderson
and Williams 2001) (Salman, 2017)		 4 0 not gregarious (inferred from photo and video material) 1 0 1 1 "spawning aggregations in space and/or time
described in many squids (Moltschaniwskyj and Pecl,
2006 and references therein), including I. coindetii
(S  anchez et al., 1998; Lefkaditou et al., 2008)." (Puerta et al., 2016)		 (Puerta et al., 2016) 0 No… "This gelatinous mass functions as a buoyancy mechanism which prevents eggs from sinking; the equilibration of density between egg masses and sea water, in fact, requires many days under most conditions. Therefore, if spawning occurs in the pelagic domain, the egg masses can remain suspended in the mesopelagic zone for a relatively long time. Such a mechanism would allow pelagically-spawned eggs to take advantage of temperature and other oceanographic conditions most beneficial for embryonic development. Illex egg masses have not been recorded in the natural environment, but observations in the laboratory of I. illecebrosus, showed that this species produces gelatinous egg masses while swimming in open water." (Jereb & Roper, 2010) (Jereb & Roper, 2010) 0 No… "This gelatinous mass functions as a buoyancy mechanism which prevents eggs from sinking; the equilibration of density between egg masses and sea water, in fact, requires many days under most conditions. Therefore, if spawning occurs in the pelagic domain, the egg masses can remain suspended in the mesopelagic zone for a relatively long time. Such a mechanism would allow pelagically-spawned eggs to take advantage of temperature and other oceanographic conditions most beneficial for embryonic development. Illex egg masses have not been recorded in the natural environment, but observations in the laboratory of I. illecebrosus, showed that this species produces gelatinous egg masses while swimming in open water." (Jereb & Roper, 2010) (Jereb & Roper, 2010) 143 161 447.888 197.532 164 80 W-W
Illex illecebrosus Illex illecebrosus 2510 "I. illecebrosus concurrently inhabits the continental shelf, slope and oceanic waters during portions of the year. However, sampling beyond the depth limit of USA and CA spring and fall bottom trawl surveys (about 366 m) is limited. Small quantities of shortfin squid have been caught in Northeast Fisheries Science Center (NEFSC) bottom trawl surveys (Azarovitz, 1981) on the upper slope between the Gulf of Maine and Cape Hatteras in April (381–460 m depths). Concurrent with the USA fishery on the continental shelf, shortfin squid were also caught offshore in July near the Bear Seamount, with the maximum catch at 2510 m depth (NEFSC, 2003), and during late fall, catch rates declined with depth and bottom temperature at depths ranging from 384 to 1,038 m, between Georges Bank and Cape Canaveral, Florida (Rathjen, 1981). Concurrent with the July inshore jig fishery off Newfoundland, shortfin squid were consistently caught offshore in Division 3M (Figure 4) bottom trawl surveys of the Flemish Cap (i.e., 20–5,143 t during 2003–2012), at depths up to 1460 m (Hendrickson and Showell, 2013)." (Arkhipkin et al., 2015) (Arkhipkin et al, 2015; Jereb & Roper, 2010; Brozniak & Hendrickson 1999; Rowell et al. 1985; Voss & Brakoniecki 1985; Vecchione & Roper, 1991, p. 434; Vecchione & Roper, 1991, p. 436; Jereb et al., 2015; Jereb et al., 2015; Roper & Young, 1975; Brozniak & Hendrickson 1999) 18 "Although common in nearshore waters north of the Gulf of
Maine during summer and fall, the species is uncommon in
shallow waters ( < 18 m) on the USA shelf (Hendrickson and
Holmes, 2004)." (Arkhipkin et al, 2015)		 (Arkhipkin et al, 2015; Jereb & Roper, 2010; Brozniak & Hendrickson 1999; Rowell et al. 1985; Voss & Brakoniecki 1985; Vecchione & Roper, 1991, p. 434; Vecchione & Roper, 1991, p. 436; Jereb et al., 2015; Jereb et al., 2015; Roper & Young, 1975; Brozniak & Hendrickson 1999) 2 **edge case** *JM Feb 28: definitely pelagic lifestyle. "We now know that Illex illecebrosus commonly rests on the sediment surface on the continental slope" (Vecchione & Roper, 1991, p. 442). "Illex illecebrosus is a pelagic Ommastrephid species, hatched at sea, coming near shore in schools as juveniles and returning to the deep ocean to spawn (O'Dor, 1983)" (Mather & O'Dor 1984). "Migrate to bottom or deeper waters during daytime. Change color to camouflaged pattern when resting on bottom to reduce risk of predation by benthic species." (Hendrickson & Holmes 2004). NW Atlantic "Most of the squid were observed near bottom at depths of about 200–900 m. The deepest depths observed were during daytime and the shallowest during the evening. However, individuals were observed as much as 1000 m above the bottom during daytime whereas others were on or near the bottom in deep water during the evening." (Harrop et al. 2014). "Illex is a genus in the family Ommastrephidae, which comprise generally pelagic squids. However, observations from submersibles have demonstrated that I. illecebrosus is very often encountered sitting motionless on the bottom (Vecchione et al. 1998) (Vecchione 2001) Mather & O'Dor 1984; Vecchione & Roper, 1991, p. 442; Hendrickson & Holmes 2004; Harrop et al. 2014 41 66 " Northwestern North Atlantic Ocean off the east coast of North America from about 26°N to 29°N off the east coast of Florida to 66°N (Iceland, southern Greenland, Baffin Island)" (Jereb & Roper, 2010) (Jereb & Roper, 2010) 25 Northwest Atlantic 25 N to 65 N (Arkhipkin et al., 2015) (Arkhipkin et al., 2015) 2 1 "The timing of migrations into the fishing areas varies interannually (Fedulov and Amaratunga, 1981) and begins earliest in the southern portion of the species’ range. During March and April, on-shelf migration occurs simultaneously along the USA shelf/slope edge, from South CaroliNAto Browns Bank on the southern Scotian Shelf, and squid densities are highest in the southernmost and deepest survey strata as well as on Browns Bank (Hendrickson, 2004). Migration onto the Scotian Shelf also begins by April (Fedulov and Amaratunga, 1981; Black et al., 1987), but migration onto the Grand Banks gener-ally occurs later, during May and June (Squires, 1957), anddensities are highest along the Bank edge in Northwest Atlantic Fisheries Organization (NAFO) Divisions 3O and 3N(Figure 4; Black et al., 1987; Hendrickson, 2006). During late May, both juveniles with a modal mantle length (ML) of 40 mm and adults were caught near the USA shelf edge (Hendrickson, 2004). By July, the species is broadly distributed across the USA shelf, Scotian Shelf and Gulf of St. Lawrence (Hendrickson, 2004, Black et al., 1987) and has migrated to the inshore fishing grounds off Newfoundland (Dawe, 1981). Fall offshore migrations also begin earliest in the southern portion of the species’ range. During September and October, squid remain distributed throughout the USA shelf but density and squid body size increase with depth for individuals greater than 100 mm ML (Brodziak and Hendrickson, 1999), indicating an off-shelf migration along the entire length of the USA shelf (Hendrickson, 2004). However, migration from the Newfoundland inshore fishing grounds occurs later, generally during November (Dawe, 1981)" (Arkhipkin et al., 2015) (Arkhipkin et al., 2015; Jereb & Roper, 2010; Luckhurst 2018; Shea et al. 2017; O'Dor & Dawe 1998; Dawe & Brodziak 1998; Perez & O’Dor 1998; Dawe et al. 1981; Sauer et al 2000; Arkhipkin 2013; Boyle, 1983; Jereb et al., 2015; 1 "The timing of migrations into the fishing areas varies interannually (Fedulov and Amaratunga, 1981) and begins earliest in the southern portion of the species’ range. During March and April, on-shelf migration occurs simultaneously along the USA shelf/slope edge, from South CaroliNAto Browns Bank on the southern Scotian Shelf, and squid densities are highest in the southernmost and deepest survey strata as well as on Browns Bank (Hendrickson, 2004). Migration onto the Scotian Shelf also begins by April (Fedulov and Amaratunga, 1981; Black et al., 1987), but migration onto the Grand Banks gener-ally occurs later, during May and June (Squires, 1957), anddensities are highest along the Bank edge in Northwest Atlantic Fisheries Organization (NAFO) Divisions 3O and 3N(Figure 4; Black et al., 1987; Hendrickson, 2006). During late May, both juveniles with a modal mantle length (ML) of 40 mm and adults were caught near the USA shelf edge (Hendrickson, 2004). By July, the species is broadly distributed across the USA shelf, Scotian Shelf and Gulf of St. Lawrence (Hendrickson, 2004, Black et al., 1987) and has migrated to the inshore fishing grounds off Newfoundland (Dawe, 1981). Fall offshore migrations also begin earliest in the southern portion of the species’ range. During September and October, squid remain distributed throughout the USA shelf but density and squid body size increase with depth for individuals greater than 100 mm ML (Brodziak and Hendrickson, 1999), indicating an off-shelf migration along the entire length of the USA shelf (Hendrickson, 2004). However, migration from the Newfoundland inshore fishing grounds occurs later, generally during November (Dawe, 1981)" (Arkhipkin et al., 2015) (Arkhipkin et al., 2015) (Roper & Young 1975; Boyle & Rodhouse, 2005) 1 “…nearly all captures occur during the daytime indicating that specimens leave the bottom and disperse into midwater at night (C. C. Lu, pers. comm.)” (Roper & Young 1975). (Roper & Young 1975) 365.3 “results indicated that the largest squid in each population sampled were no more than about 1 year old. (Hurley and Beck,
1979; Radtke, 1983)” (Rodhouse and Hatfield, 1990)		 (Coelho et al. 1994; Arkhipkin & Fetisov 2000; Perez & O’Dor 2000; Ragonese et al. 2002; Hendriskson 2004; Hendrickson & Hart 2006; Arkhipkin et al., 2015; Rodhouse and Hatfield, 1990) 115 "The lifespan of mated females from the winter cohort inhabiting the USA shelf was 115–215 days (Hendrickson, 2004) whereas a maximum age of 250 days was documented for females caught in the Newfoundland jig fishery and which were not mature (Dawe and Beck, 1997). The species exhibits latitudinal clines in growth rate and size-at-maturity such that individuals inhabiting warmer waters of the Mid-Atlantic Bight exhibit faster growth and maturation rates, and possibly have a shorter lifespan, than squid from the colder waters off Newfoundland (Hendrickson, 2004)." (Arkhipkin et al., 2015) (Coelho et al. 1994; Arkhipkin & Fetisov 2000; Perez & O’Dor 2000; Ragonese et al. 2002; Hendriskson 2004; Hendrickson & Hart 2006; Arkhipkin et al., 2015; Rodhouse and Hatfield, 1990) 92 "Minimum age-at-maturity was 88 d for males and 92 d for females. Females attained 50% maturity at a younger age (A50 ¼
144 d) than males (A50 ¼ 154 d), but this difference was not
significant"(Hendrickson 2004) 		(Hendrickson 2004) 88 "Minimum age-at-maturity was 88 d for males and 92 d for females. Females attained 50% maturity at a younger age (A50 ¼
144 d) than males (A50 ¼ 154 d), but this difference was not
significant"(Hendrickson 2004) 		(Hendrickson 2004) 7 6 0 "In Illex illecebrosus, too, Nicol and 0'Dor {I985} observed head-first attacks on swarms of the krill Meganyctiphanes norvegica in the Bay of Fundy [i.e., cooperative hunting?], but noted that individuals in a small shoal captured the euphausiids repeatedly but independently of each other, attacking the body of the swarm rather than peripheral members and showing no signs of cooperative hunting. Illex behave similarly in the laboratory [Foyle & O'Dor, 1987)." (Hanlon & Messenger, 2018, p. 85) Hanlon & Messenger, 2018, p. 85 1 "A good example of stalking is the attack of the squid Illex illecebrosus on trout (Fig. 4.9). This involves a head-first attack after a long tracking phase. in which the squid gradually closes in on the trout from behind, where its visibility is poorest. Interestingly, this particular behaviour is reserved for large, fast prey: smaller, slower fish are caught without stalking, by tail-first attacks. Hanlon & Messenger, 2018, p. 83) Hanlon & Messenger, 2018, p. 83)
(Foyle & O'Dor 1988
Jastrebsky et al 2017		 1 When swarms of pelagic crustaceans euphausids, for example, are encountered, the squid rapidly flares and expands its arms to create an in-flowing trbulence in which to trap, then encircle the prey." (Jereb & Roper, 2010) Jereb & Roper, 2010 1 No tracking phase occurs in attacks on mummichogs (small prey). Fishes are captured by a quick headfirst atack with rapidly out-thrust tentacles, which then withdraw the fish into the open arm crown; the entire process is accomplished in less than 2 seconds. (Jereb & Roper, 2010) Jereb & Roper, 2010
Nicol & O’Dor 1984
Squires 1966
Kier 1991		 1 When swarms of pelagic crustaceans euphausids, for example, are encountered, the squid rapidly flares and expands its arms to create an in-flowing trbulence in which to trap, then encircle the prey." (Jereb & Roper, 2010) Jereb & Roper, 2010 1 "squid bit the fish, severing their spinal cord as a complementary predation behaviour, securing prey of high risk of escape. Similar hunting strategy has also been observed for Illex illecebrosus feeding on young mackerel (Verrill 1882)" (Carreno Castilla et al 2020) Carreno Castilla et al 2020
Bradbury & Aldrich 1969		 1 After filleting the fish, the squid dropped the head and spinal column which sank to the bottom of the tank. (Foyle & O'Dor 1988) Foyle & O'Dor 1988 1 These differences in hunting tactics can be explained in terms of the mechanical limitations of the squid's jet propulsion system: head-first acceleration is poor in Illex compared with the trout, which could not be caught unless the squid got close to it (Foyle & O'Dor, 1987). Illex also attack jigs head first (Williamson, 1965)." (Hanlon & Messenger, 2018, p. 83) Hanlon & Messenger, 2018, p. 83)
Jereb and Roper 2010
(Foyle & O'Dor 1988
Uchikawa and Kidokoro, 2014		 5 4 1 "illecebrosus was observed both in midwater and on the bottom, where it rests on its arm tips with its head, mantle opening and funnel raised off the substrate. Resting sites apparently are selected because of their paucity of epibenthic fauNAas potential predators." (Jereb & Roper, 2010) Jereb & Roper, 2010
O'Dor & Dawe 1998		 1 “When attempts were made to remove squid from the circular tanks for various reasons, a colouration change, from dark to complete translucency, was noted and interpreted as being of escape value." (Bradbury 1974) Bradbury 1974 1 Two types of inking were observed. In the first, most often observed when squid were taken on jiggers in the field and placed in suitable containers for transportation, a dilute seawater solution of ink was emitted. Such ink was observed to be rapidly dispersed in seawater and caused a mere cloudy discolouration of the water. The second type is more viscous, forming strings of inky black water mixed with what appears to be mucus. This ink, upon emission, remained in discrete "shapes" in the water for longer periods of time than did the dilute ink of the other type…This latter type of inking occurred when attempting to remove squid from a maintenance tank in the Laboratory. Both types of inking were accompanied by violent contractions of the mantle musculature, with subsequent elimination of sea water from the mantle cavity.” (Bradbury 1974) Bradbury 1974 1 "squid [i.e., Illex illecebrosus] that had been singled out from their shoal by dolphins survived attack only if they used the tactic of changing direction erratically as they also changed body patterns and inked (Major, 1986)." (Hanlon & Messenger, 2018, p. 129) Hanlon & Messenger, 2018, p. 129 1 Besides the "tactics" of schooling and hiding under the vessel, individual squid, closely «0.5 m) pursued by individual dolphin, released a ball or cloud of ink, simultaneously changing direction. In one-on-one interactions, high-speed unpredictable changes in direction occurred as each squid attempted evasion with or without the release of ink. In three interactions observed in their entirety just under the surface in the well-illuminated area, the entire chase was almost linear in trajectory and led to the successful capture of a squid by a dolphin. In each instance, the chase covered approximately 5 m, the squid having been isolated from a school by the dolphin. The squid's color appeared to change from a dark (red) to light as it moved into the center of the illuminated field. As the dolphin decreased its distance to the squid to about 0.5 m, the squid released a small cloud of ink, which the dolphin swam through; the ink passing across the rostrum and eyes of the dolphin. The squid continued to move horizontally, then stopped to be overrun and consumed, tentacles first, by the dolphin. Observation of individual squid releasing ink and escaping after changing direction abruptly provides some evidence that dolphins swimming through the ink cloud may have been using visual and not acoustic means to track the squid. Debilitated or disoriented squid were not observed. Such squid might have provided some evidence that intense sound pulses were being used by the dolphins to stun their prey as has been hypothesized by Norris and Mohl (1983). The observations reported here closely parallel similar observations made of interactions between visual orienting schooled and individual predatory fishes and schooled and individual prey fishes (Major, 1977, 1978).” (Major 1986) Major 1986 40 Jereb & Roper, 2010 O'Dor & Dawe 1998
Jereb & Roper 2010
Dawe & Brodziak 1998
Dawe et al. 1997
Lange & Sissenwine 1980
O'Dor 1983
Xavier et al 2018
Boyle 1983		 0 0 5 35 "The
predators of I. illecebrosus are numerous. Certain size ranges of I.
illecebrosus are most vulnerable to predation, and predation on the squid
varies ontogentically among predators such as finfishes, elasmobranchs and
marine mammals. Illex illecebrosus is a significant prey of the cod fish (Gadus
morhua), swordfish (Xiphias gladius), other billfishes and tunas, bluefish
(Pomatomus saltatrix), goosefish (Lophius americanus), silver hake (Merluccius
bilinearis), summer flounder (Paralichthys dentatus), shortfin mako shark
(Isurus oxyrinchus) and bigeye thresher shark (Alopias superciliosus); seabird
predators include: northern gannet, Murus bassanus, greater shearwaters
(Puffinus gravis), sooty shearwaters (P. griseus), fulmars (Tulmarus
glacialis), and the larger gulls, e.g. black-backs.

Illex illecebrosus was the nearly exclusive prey of the long-finned pilot whale
(Globicephala malaena) in Newfoundland inshore waters into the mid-1970s (up to
10 million tonnes annually); their distributions and seasonal movements
coincide in time and space. Peaks in landings were coincident and the
availability of pilot whales in shore was dependant on I. illecebrosus
abundance. Annual squid production (biomass), based on consumption of I.
illecebrosus by pilot whales, was in the order of several hundred thousand
tonnes.

Other mammals predators include the white-sided dolphins (Lagenorhychus
acutus), the “squid hound” (L. albirostris) and the harbor seals (Phoca
vitulina). As a consequence, the international fishery for I. illecebrosus
captures marine mammals as incidental catch; pilot whale species (Gloubicephala
spp.) and common dolphin (Delphinus delphis), together comprising 93% of
incidental catch in Canadian and the United States waters, southward into the
mid-Atlantic Bight. These species are known to be major predators on I.
illecebrosus, so quite naturally are foraging in the squid concentrations
during trawling operations. "
(Jereb & Roper, 2010)

 (Jereb
& Roper, 2010) (Staudinger
et al. 2012) (Shea
et al. 2017) (Dawe
& Brodziak 1998) (Dawe
& Mercer 1982) (Xavier
et al., 2018) (Craddock
et al 2009) (Boyle,
1983) Hanlon
& Messenger, 2018, (Staudinger
et al. 2013)		 4 3 shoaling/schooling “Illex illecebrosus squid appear to have a species-typical and internally organized spatial arrangement of their groups. Squid maintained an average angle of 25° with respect to their nearest neighbour, and mostly had angular deviations between 5° and 20°. They maintained distances to nearest, second and third neighbours in a ratio of 1:1.5:2. The distances were strongly affected by group size (4, 20, or 38), with larger groups maintaining closer distances." (Mather & O'Dor 1984) Major (1986), cited in Hanlon & Messenger, 2018, p. 129.; Mather & O'Dor 1984 1 shoaling/schooling “Illex illecebrosus squid appear to have a species-typical and internally organized spatial arrangement of their groups. Squid maintained an average angle of 25° with respect to their nearest neighbour, and mostly had angular deviations between 5° and 20°. They maintained distances to nearest, second and third neighbours in a ratio of 1:1.5:2. The distances were strongly affected by group size (4, 20, or 38), with larger groups maintaining closer distances." (Mather & O'Dor 1984) Major (1986), cited in Hanlon & Messenger, 2018, p. 129.; Mather & O'Dor 1984 4 1 1 "It is probably not always possible for a squid to swim past a small active fish sufficiently close or fast to capture it in the turbulent wake set up by expanding its arms. Observations of the capture of fish by Illex reveal that although the squid swims past the fish or lure, it then turns and darts with arms forward and grabs its prey. This invariably causes the schooling pattern to break up. It is quite probable that there is a connection between break-up of schools and the not uncommon finding of squids in stomachs of inshore squids. The change from feeding mainly on small crustaceans to feeding mostly on fish may in part be responsible for this. It would seem that squids have to learn this method of feeding because in the first seasonal occurrence of squid in the inshore zone there is a period in which fish-shaped lures do not attract them (the period during which fishermen consider the squids to be "blind")." (Squires 1966) Squires 1966 3 "The population dynamics of I. illecebrosus and the strategies employed to maintain them are quite complex and may include kinship, school cohesion and cannibalism" (Jereb & Roper, 2010) Jereb & Roper, 2010 1 “Illex illecebrosus squid appear to have a species-typical and internally organized spatial arrangement of their groups. Squid maintained an average angle of 25° with respect to their nearest neighbour, and mostly had angular deviations between 5° and 20°. They maintained distances to nearest, second and third neighbours in a ratio of 1:1.5:2. The distances were strongly affected by group size (4, 20, or 38), with larger groups maintaining closer distances." “Casual observations showed that squid schooled much more closely after visual threat such as the presence of a dip net or figures on the edge of the pool. Individual squid were seldom away from the school for more than a minute, and quickly moved back to the school if artificially separated from it. When a school was divided by dropping a large 4 mm mesh seine net across the centre of the pool, the two halves of the school initially swam back and forth together near the net rather than avoiding it. This behaviour "decayed" and the groups separated after they became accustomed to the net. The normal response to the same net when the school is not divided is avoidance. When the squid were fed either small fish singly or several fish, some individuals appeared to have prior access to food despite proximity, catching several and appearing to "dominate" other individuals…Casual observations at feeding time suggested the possibility of a dominance hierarchy, with some individuals capturing several fish and making "rushes" at others which prevented them feeding” (Mather & O'Dor 1984) & "At times, several squid would converge on the introduced capelin and engage one another in conflict in an effort to secure the prey (Fig. 4H)… It was observed that the contesting squid alternated between a dark and a light coloration during the conflict, with squid “’b” blanching considerably before disengagement (Figs, 4F and 4G). Both squid were of approximately the same size (24 +/- 2 cm in mantle length) and we are unable to assign any cause for one to be the victor” (Bradbury & Aldrich 1969) (Mather & O'Dor 1984) 1 "The only confirmed spawning area is located along the USA shelf edge, in the Mid-Atlantic Bight (between 39 10 ´ N and 35 50 ´ N), where the winter cohort was found spawning during late May at depths of 113–377 m and surface and bottom temperatures ranging from 13.4–20.1 C and 11.4–20.3 C, respectively (Hendrickson, 2004). Mature and spawning individuals have also been caught in the USA directed bottom trawl fishery during June-September (Hendrickson and Hart, 2006). Thus, the Mid-Atlantic Bight is the primary spawning area during at least May–September, but some spawning may also occur in the Gulf Stream/Slope Water frontal zone where paralarvae and juveniles have been collected during most winter months (Hatanaka et al., 1985b). The presence of spawners during May–September, combined with the documentation of November–June hatch dates (Dawe and Beck, 1997; Hendrickson, 2004), indicate that spawning occurs year-round." (Arkhipkin et al., 2015) (Arkhipkin et al., 2015) 1 “The sex ratio may be a determinant of spawning success and fecundity because males appear to be the initiators of mating behaviour ( O ‘ Dor et al . , 1980b ). Figure 11.7 summarizes an experiment in which two schools of squid were held in the 15 m pool for several months. The pattern observed suggests that when a male is ‘fully mature’ ( precisely what this means is unclear ) it will mate with several females. With only one exception mated females were vitellogenic (stage IV) and with only two exceptions they had oviducal eggs present (late state IV or stage V) . This selectivity may reflect an element of female receptiveness since some females resist. Only one or two brief episodes of courtship behaviours of the sort described for loliginids ( Drew , 1911 ; Arnold , 1962 ; Fields , 1965 ) have been observed despite extensive efforts , suggesting that courtship is either less pronounced or more secretive in Illex illecebrosus” (O’Dor 1983) (O’Dor 1983) 0 "Eggs enclosed in neutrally buoyant gel masses are carried north by the Gulf Stream, where embryonic development and hatching takes place. The early life is spent along the meandering northern boundary of the Gulf Stream and the slope waters, and it ends when the offspring reach the adult habitat on the continental shelf. During this transition, offspring are subject to important temperature and food availability gradients, as determined by their encountering distinct water masses. The squid being provided with limited yolk supply and an energetically expensive life-style (O'Dor et al. 1986), such gradients may impose serious limitations on the attainment of required growth rates and survival during the early life.” (Perez & O’Dor 1998) (Perez & O’Dor 1998) 0 "Eggs enclosed in neutrally buoyant gel masses are carried north by the Gulf Stream, where embryonic development and hatching takes place. The early life is spent along the meandering northern boundary of the Gulf Stream and the slope waters, and it ends when the offspring reach the adult habitat on the continental shelf. During this transition, offspring are subject to important temperature and food availability gradients, as determined by their encountering distinct water masses. The squid being provided with limited yolk supply and an energetically expensive life-style (O'Dor et al. 1986), such gradients may impose serious limitations on the attainment of required growth rates and survival during the early life.” (Perez & O’Dor 1998) (Perez & O’Dor 1998) 251 160 198.4 162 NY
Japetella diaphana Japetella diaphana 2500 "[Thore, 1949] also reported ontogenetic descent in this species; the juveniles (25 mm) were found between 100-300 m while the adults were concentrated between 1750-2500 m." (Cairns, 1976) (Cairns, 1976; Judkins & Vecchione, 2020; Jereb et al., 2014; Urbano & Hendrickx, 2018; Birk et al., 2019; ) 600 [AL: Caught closer to the surface but no clear evidence for adults] One hundred fifty-three Japetella diaphana (5–92 mm ML)
follow a similar pattern, with no obvious diel migration pattern,
and those larger than 26 mm ML living below 600 m, with
one exception where a 44 mm ML specimen was collected
between 0 and 200 m" (Judkins & Vecchione, 2020)		 (Cairns, 1976; Judkins & Vecchione, 2020; Jereb et al., 2014; Urbano & Hendrickx, 2018; Birk et al., 2019; ) 1 "pelagic" (Jereb et al., 2014) Jereb et al., 2014 100 60 60 N to 40 S (Jereb et al. 2014) (Jereb et al. 2014) -40 60 N to 40 S (Jereb et al. 2014) (Jereb et al. 2014) 3 1 "[Thore, 1949] also reported ontogenetic descent in this species; the juveniles (25 mm) were found between 100-300 m while the adults were concentrated between 1750-2500 m." (Cairns, 1976) (Cairns, 1976; Jereb et al., 2014; Romero et al., 2020; Roper & Young 1975) NA 0 1 1 1 "They are gelatinous and can hide from predators by rapidly changing between being transparent or pigmented depending on the predatory search strategy (Zylinski and Johnsen, 2011)" (Birk et al 2019) Birk et al 2019 6 "Analysis of the digestive tracts of specimens of up to 20 mm ML showed the presence of calanoid, euphausiid, and decapod crustaceans, molluscs, chaetognaths, and fishes in their diet (Passarella and Hopkins 1991)." (Nixon & Young 2003) Nixon and Young 2003
Passarella and Hopkins		 0 1 Nixon and Young 2003 4 7 "Cuvier’s
beaked whale; blue shark; pelagic osteichthyes; Ommastrephes bartramii"
(Xavier et al., 2018)		 (Zambrano-Zambrano et al., 2019) (Vaske
Júnior et al. 2009) (Xavier
et al., 2018) (Tscuchiya
et al. 1998) (Tsuchiya
and Sawadaishi 1997)		 4 0 not gregarious (inferred from photo and video material) 2 0 2 "These females
possess a large yellow photophore around their mouth, one of the few
known bioluminescent structures in any octopod, presumably to attract
males (Robison and Young, 1981; Herring et al., 1987)." (Birk et al 2019)		 "Nearly mature males have salivary glands that are much larger than those of comparable females. As in Bolitaena, salivary products may be used as a chemical attractant for females. The female light organ may be used for reproductive signalling to males" (Jereb et al., 2014) 14 13 5.2 25 NY
Joubiniteuthis portieri Joubiniteuthis portieri 4000 one specimen found in Nova Scotia minimum depth 4000m (Vecchione & Pohle 2002) (Jereb & Roper, 2010; Vecchione & Pohle 2002; Judkins & Vecchione, 2020) 0 "Nineteen J. portieri (5–159 mm ML) documented the
species living throughout the water column; six were found in day tows inhabiting the upper and lower mesopelagic
zone while thirteen individuals were collected at night
from the surface to 1500 m. There is some evidence of
vertical migration but no ontogenic shift for this species" (Judkins & Vecchione, 2020)		 (Jereb & Roper, 2010; Vecchione & Pohle 2002; Judkins & Vecchione, 2020) 1 "living throughout the water column" (Judkins & Vecchione, 2020) (Judkins & Vecchione, 2020 70 40 "Cosmopolitan in tropical and subtropical, even temperate waters; Atlantic Ocean, 40°N to 30°S,
including Caribbean Sea; Pacific Ocean, Hawaii, Japan, eastern Australia, Tasman Sea, New Zealand (Fig. 254)." (Jereb & Roper, 2010)		 (Jereb & Roper, 2010) -30 "Cosmopolitan in tropical and subtropical, even temperate waters; Atlantic Ocean, 40°N to 30°S,
including
Caribbean Sea; Pacific Ocean, Hawaii, Japan, eastern Australia, Tasman
Sea, New Zealand (Fig. 254)." (Jereb & Roper, 2010)		 (Jereb & Roper, 2010) 4 0 "no [evidence of] ontogenic shift for this species" (Judkins & Vecchione, 2020) (Judkins & Vecchione, 2020) 0 "no [evidence of] ontogenic shift for this species" (Judkins & Vecchione, 2020) (Judkins & Vecchione, 2020) NA 0 NA 0 0 0 3 "Specimens
are reported as prey of lancetfish, blue shark and sperm whale." (Jereb
& Roper, 2010)		 Jereb & Roper, 2010 3 0 not gregarious (inferred from photo and video material) NA 0 NA 4 5 7 36 NY
Leachia dislocata Leachia atlantica 1879 Caught from 865 m to 1879 m (Urbano & Hendrickx, 2019) (Jereb & Roper, 2010; Urbano & Hendrickx, 2019; Urbano and Hendrickx, 2018) 200 "Juveniles occur in the upper few hundred metres of the epipelagic zone; then with ontogenetic growth they descend into the bathypelagic depths of 1 000 m or more. Mating and spawning appear to occur in the upper 200 m, based on captures of mature males and a spent female in those upper epipelagic waters." (Jereb & Roper, 2010) (Jereb & Roper, 2010; Urbano & Hendrickx, 2019; Urbano and Hendrickx, 2018) 1 Pelagic (Urbano & Hendrickx, 2019) Urbano & Hendrickx, 2019 83 45 "This species occurs in oceanic waters of the eastern North Pacific Ocean in the California Current
between approximately 25°N and 45°N and westward in North Pacific central waters to 160°W, including the Hawaiian Islands;
in the Peru-Chile Current between about 15°S and 25°S. " (Jereb & Roper, 2010)		 (Jereb & Roper, 2010) -38 ""This species has been found on both sides of the equator in the Pacific and Atlantic Oceans, and in the Indian Ocean…In the New Zealand region L. eschscholtzii has been taken between 28°S and 38°S" (Imber 1978) (Imber 1978) 3 1 "Juveniles occur in the upper few hundred metres of the epipelagic zone; then with ontogenetic growth they descend into the bathypelagic depths of 1 000 m or more. Mating and spawning appear to occur in the upper 200 m, based on captures of mature males and a spent female in those upper epipelagic waters." (Jereb & Roper, 2010) (Jereb & Roper, 2010) NA 0 NA 0 0 0 2 "An opportunistic surface feeding event was recorded during the night of 7 October,
2007. A mature female (visually estimated at 6m long) was seen very close to
shore, in the vicinity of Primer Cañon, swimming slowly at the surface with the
mouth open to feed on a concentration of squid identified as Leachia dislocata
(Family Cranchiidae; Figure 4). Some squid were dead and floating on the
surface so likely the Cuvier’s beaked whale individual took most of the squid
by scavenging. The female approached the boat as near as one meter away and
after a near inspection it moved to the north slowly. According to MacLeod et
al. (2003), this family of squid was one of the most frequently reported on
dietary data available for 38 specimens of Cuvier’s beaked whales from
throughout the range of the species. This squid has also been reported as prey
of the Laysan albatross (Phoebastria immutabilis) at Guadalupe Island (Pitman
et al., 2004). This seabird is a daytime sea surface scavenger of spawned adult
squid (Pitman et al., 2004). L. dislocata are available to albatrosses because,
as in many squid species, females die after spawning and float to the surface
(Nesis et al., 1988)" (Cardenas-Hinojosa et al., 2015)		 (Cardenas-Hinojosa
et al., 2015)		 2 1 sp unspecified "Capture records of near mature females show them to be solitary, with an estimated minimum separation of 61 m. The density of the females is within a reasonable order of magnitude for sexual attraction by bioluminescence to be feasible, especially in depths of l000m or more where background bioluminescence is low and predators are scarce (R.E. Young 1983)." (Nixon & Young 2003) Nixon & Young 2003 0 sp unspecified "Capture records of near mature females show them to be solitary, with an estimated minimum separation of 61 m. The density of the females is within a reasonable order of magnitude for sexual attraction by bioluminescence to be feasible, especially in depths of l000m or more where background bioluminescence is low and predators are scarce (R.E. Young 1983)." (Nixon & Young 2003) Nixon & Young 2003 NA 0 NA 4 4 17.8 132 NY
Loligo forbesii Loligo forbesii 1000 "It occurs over the continental shelf in the temperate part of its distributional range, but it is found in deeper waters in subtropical areas. Its entire vertical range extends from depths shallower than 50 to over 700 m, while the Azores population occurs deeper than 1 000 m. In the Mediterranean Sea, it very seldom occurs in waters less that 80 to100 m depth, and its bathymetric range overlaps only slightly with that of its congener Loligo vulgaris. In Atlantic waters depth distribution varies by season, with squid remaining mostly in waters along the shelf edge (100 to 200 m), then gathering in waters of less than 50 m during the spawning peaks" (Jereb & Roper, 2010) (Jereb & Roper, 2010; Arkhipkin et al., 2015; Quetglas et al., 2000; Sifner et al., 2005; Silva et al., 2011; D’Onghia et al. 2011; Valls et al 2015; Oesterwind et al 2010; Morris et al., 1993; Moreno et al 1994; Jereb et al., 2015; Barrett et al 2021; Vafidis et al 2008) 5 "depth 5-346 m" (Vafidis et al 2008) (Jereb & Roper, 2010; Arkhipkin et al., 2015; Quetglas et al., 2000; Sifner et al., 2005; Silva et al., 2011; D’Onghia et al. 2011; Valls et al 2015; Oesterwind et al 2010; Morris et al., 1993; Moreno et al 1994; Jereb et al., 2015; Barrett et al 2021; Vafidis et al 2008) 2 **edge case** "In Scotland, a small directed fishery takes L. forbesii in inshore waters over rocky ground at the start of the season, and over sandy/muddy bottom later in the season (Young et al., 2006; Smith, 2011), possibly indicating a shift in habitat preference with maturity" (Smith et al 2013). "demersal…fatty-acid and stable-isotope analyses have shown the association between L. forbesii and the benthic food web" (Rosa et al. 2013) Smith et al 2013;
Jereb & Roper, 2010		 43 63 "L. forbesi occurring
throughout the northeast Atlantic between 20 and 63◦N" (Chen et al 2006)		 Chen et al 2006) 20 60 N (about 64 N on the map) to 20 N (Jereb & Roper, 2010) (Jereb & Roper, 2010) 1 1 "It shows an onshore-offshore ontogenetic migration, typical of loliginids, moving from the shelf edge (at 100–200 m) in summer toward inshore waters to spawn in the winter. In some years a West-East migration apparently occurs in autumn in Scottish waters (Waluda and Pierce, 1998). In the Moray Firth, Scotland, the smallest individuals are caught close inshore in summer and there seems to be a subsequent ontogenetic migration away from the coast and a later return of mature animals into coastal waters to spawn (e.g., ViaNAet al., 2009)." (Arkhipkin et al., 2015) (Jereb & Roper, 2010; Arkhipkin et al., 2015) 570 “The relationship between age and sexual maturity of male and females in both species was weak (Fig. 3). Immature and maturing specimens were present at a wide range of ages in both sexes and species, especially in males of L. forbesii…L. forbesii became mature later than L. vulgaris; with females becoming mature from 399 days on, and the females in the 450 age class were always mature. Males could be mature from 310 days, but immature and maturing specimens were still abundant in the 480–540 age classes; only one specimen was found to be mature at the 570 age class (506.0 mm ML and 1910.0 g)” (Agus et al. 2018) (Agus et al. 2018; Hanlon et al. 1989) 249 “The smallest mature male with spermatophores in the penis was 102 mm ML at day 249 (LF83). Some females began to produce maturing ova at about 90-l00mm ML (Fig. 7), corresponding to 250-390 days in culture (Fig. 6 and Forsythe & Hanlon 1989).” (Hanlon et al. 1989) (Agus et al. 2018; Hanlon et al. 1989) 2 2 1 lab "appearance of the chromatic component Dark first arms" when seizing prey (Porteiro et al. 1990) Porteiro et al. 1990 1 . The behaviour we observed during seizure and head cutting when the prey was less than about 35 cm, follows in general the description given by Bidder (1950) for L. vulgaris. However, we draw attention to the preference in using the first and second arms in manipulating the prey and the appearance of the chromatic component Dark first arms. (Porteiro et al. 1990) Porteiro et al. 1990 NA 0 65 "The veined squid is a highly opportunistic predator, feeding on fish, crustaceans, cephalopods, polychaetes, and any other potential available prey, and cannibalism, also, occurs" (Arkhipkin et al., 2015) Arkhipkin et al., 2015
Valls et al 2015
Wangvoralak et al 2011
Martins 1982
Jereb et al 2015		 0 1 Martins 1982 7 24 "Table
12.4. Known predators of Loligo forbesii in the Mediterranean Sea and Northeast
Atlantic." (Jereb
et al., 2015)

Cephalopoda: European squid (Loligo vulgaris) Guerra and Rocha (1994)

Chondrichthyes: Blonde ray (Raja brachyura) Farias et al. (2006)

Osteichthyes: Atlantic cod (Gadus morhua) Daly et al. (2001), Magnussen (2011)

Atlantic lizardfish (Synodus saurus) Soares et al. (2003)

Monkfish (Lophius piscatorius) Daly et al. (2001)

Swordfish (Xiphias gladius) Salman (2004)

Aves: Great skua (Catharacta skua) Furness (1994)

Pinnepedia: Grey seal (Halichoerus grypus) Pierce et al. (1991a)

Harbour seal (Phoca vitulina) Brown and Pierce (1998)

Cetacea: Harbour porpoise (PhocoeNAphocoena) Rogan and Berrow (1996), Santos
et al. (2005b)

Risso's dolphin (Grampus griseus) Bearzi et al. (2011)

Sperm whale (Physeter macrocephalus) Clarke and Pascoe (1997), Santos et al.
(1999*, 2001b)

Striped dolphin (Stenella coeruleoalba) Würtz and Marrale (1993)

(Katsanevakis
et al 2008) (Jereb et al., 2015) (Bloch et al. 2012) (Pierce et al., 2010) (Velasco
et al., 2001) (Grellier and Hammond 2006) (Kousteni et al. 2018) (Arkhipkin et
al., 2015)		 5 3 "social squid" according to Hanlon & Messenger (1996:46). “Young squids attempt to form schools as soon as they are large enough to swim well against a current." (Hanlon et al. 1989) Hanlon & Messenger 1996:46; Hanlon et al. 1989 1 "social squid" according to Hanlon & Messenger (1996:46). “Young squids attempt to form schools as soon as they are large enough to swim well against a current." (Hanlon et al. 1989) Hanlon & Messenger 1996:46; Hanlon et al. 1989 3 0 3 1 "Loligo forbesi may become more numerous (or aggregated) during the spawning season (Collins et al. 1999)." (Hastie et al., 2009) (Hastie et al., 2009) 1 "Males consistently exhibit at least two alternative growth strategies, with some matur-
ing very small; these probably correspond to different reproductive strategies: mate-
guarding by large males and sneaking by small males (Hanlon and Messenger, 1998).
However, some studies have suggested as many as 3–4 cohorts (or “microcohorts”),
with different growth trajectories in males and 2–3 in females (e.g. Collins et al., 1999).
It should be noted, however, that reliable resolution of multiple modes in length fre-
quency data requires large samples and regular sampling. " (Jereb et al., 2015)		 1 "Males consistently exhibit at least two alternative growth strategies, with some matur-
ing very small; these probably correspond to different reproductive strategies: mate-
guarding by large males and sneaking by small males (Hanlon and Messenger, 1998).
However, some studies have suggested as many as 3–4 cohorts (or “microcohorts”),
with different growth trajectories in males and 2–3 in females (e.g. Collins et al., 1999).
It should be noted, however, that reliable resolution of multiple modes in length fre-
quency data requires large samples and regular sampling. " (Jereb et al., 2015)		 298 214 283.6 50 NY
Loligo vulgaris Loligo vulgaris 545 "Usually more abundant in waters shallower than 100 m, L. vulgaris is found from the coast to the limits of the upper slope (200–550 m) (Jereb et al., in press), the deepest record in the Mediterranean being at 545 m, in the eastern Ionian Sea (Krstulovi  c  Sifner et al., 2005). Where its distribution overlaps with that of L. forbesii, L. vulgaris tends to be found in shallower waters, the switch from dominance of one species to the other being placed at around 70–80 m (Ragonese and Jereb, 1986; Ria et al., 2005)." (Arkhipkin et al., 2015) (Jereb & Roper, 2010; Arkhipkin et al., 2015; Salman et al., 1998; Sifner et al., 2005; Silva et al., 2011; Roditi et al., 2020; Ruby and Knudsen 1972; Valls et al 2015; Sanchez and Demestre 2010; Mathger and Denton, 2001; Pereira et al., 1997; Adam, 1937; Sen 2005-06; Villanueva et al 2003; Moreno et al 1994; Sanchez and Guerra 1994; Jereb et al., 2015; Boyle, 1983; Bas, 1975; Roper & Young, 1975; Vafidis et al 2008) 2 In Kalymnos Island (South Aegean Sea) the species was caught with handlines in depth ranging from 2-73 m with a mean of 30.29 +/- 17.92 (Roditi et al., 2020) (Jereb & Roper, 2010; Arkhipkin et al., 2015; Salman et al., 1998; Sifner et al., 2005; Silva et al., 2011; Roditi et al., 2020; Ruby and Knudsen 1972; Valls et al 2015; Sanchez and Demestre 2010; Mathger and Denton, 2001; Pereira et al., 1997; Adam, 1937; Sen 2005-06; Villanueva et al 2003; Moreno et al 1994; Sanchez and Guerra 1994; Jereb et al., 2015; Boyle, 1983; Bas, 1975; Roper & Young, 1975; Vafidis et al 2008) 2 **edge case** "Loligo vulgaris generally has a pelagic habitus, but it becomes more dependent on the bottom during spawning seasons" (Jereb & Roper, 2010). "nectobenthic neritic species that inhabits the circumlittoral region in subtropical/temperate waters (Hastie et al., 2009; Jereb and Roper, 2010)…typically lives shallower than 100 m (Sanchez and Guerra, 1994; Salman et al., 1997; Sanchez et al., 1998; Tserpes et al., 1999) (Rosa et al. 2013). "in the lower half of the water column, may be classified as a benthopelagic species, although also as part of the shelf demersal community from a fisheries perspective" (Moreno et al. in Rosa et al.) Jereb & Roper, 2010 77 57 ""occurs along the eastern Atlantic from the British Isles (55 ° N) to the 
Gulf of Guinea (20 ° S; Roper et al., 1984), extending out to Madeira (Clarke and Lu, 
1995), and throughout the Mediterranean Sea (Belcari, 1999a). Adults are occasionally 
reported off the northwest coast of Scotland at 57°N (Pierce et al., 1994a; Hastie et al., 
2009a), the North Sea (De Heij and Baayen, 2005), and the Kattegat and western Baltic 
Sea (Jaeckel, 1937; Muus, 1959; Hornborg, 2005). Paralarvae are absent north of the 
English Channel (Yau, 1994; Collins et al., 2002). L. vulgaris is a nectobenthic species 
that lives in the circumlittoral and upper bathyal zones (Worms, 1983). " (Pierce et al., 2010)		 (Pierce et al., 2010) -20 63 N to 20 S from looking at the map, though Jereb and Roper (2010) writes " Eastern Atlantic Ocean: from approximately 55°N, around the British Isles, the North Sea (including the Skagerrak, the Kattegat and the western Baltic Sea), to 20°S, off the southwestern coast of Africa, including Madeiran waters. Mediterranean Sea: from the western to the eastern basins, including the Adriatic Sea " (Jereb & Roper, 2010) (Jereb & Roper, 2010) 1 1 "Migratory patterns have been described in the northeast Atlantic, but are still poorly understood (Jereb et al., in press)" (Arkhipkin et al., 2015) (Arkhipkin et al., 2015; Hastie et al. 2009; Paulij et al 1990;ViaNAet al. 2009; Jereb et al., 2015; Elsevier, 2014) 1 "There was evidence of a summer/autumn migration of squid
into both ends of the North Sea. An influx of squid
into the southern North Sea from the English Channel
via the Straits of Dover was recorded by Holme (1974)
for Loligo forbesi." (Waluda and Pierce 1998)		 (Bas, 1975; Roper & Young, 1975; Smith et al 2013; ) 1461 "Estimates based on length-frequency analysis suggest that L. vulgaris can live for up to
4 years (Mangold-Wirz, 1963a). However, counts of daily growth increments in stato-
liths reveal that lifespan is normally ca. 1 year, although variations have been reported,
as detailed hereafter. Slightly longer lifespans (382 and 396 d) have been recorded in
males from Galician waters (Rocha and Guerra, 1999) and the West Saharan shelf
(Arkhipkin, 1995). Maximum lifespans of 15 months in both sexes were observed in
northwestern Portuguese waters by Moreno et al. (2007). Note, however, that both Bet-
tencourt et al. (1996) and Raya et al. (1999) estimated rather shorter lifespans: 9 months
in southern Portuguese waters and 10 months on the western Saharan shelf, respec-
tively; it is not clear whether this represents real biological variation or whether meth-
odological issues are partly or wholly responsible. " (Jereb et al., 2015)		 (Moreno et al 2007; Arkhipin 1995; Rocha and Guerra 1999; Arkhipkin 1995; Jereb & Roper, 2010; Natsukari and Komine 1992; Boyle, 1983; Arkhipkin et al., 2015; Jereb et al., 2015; Pierce et al., 2010) 273.9 "Life span duration based on statolith analysis is estimated to range between 9 and 10 months (southern Portuguese and West Saharan shelf waters) and 1.5 years (north Portuguese waters), considerably shorter than the 2 to 4 years previously estimated on the basis of length frequency analyses." (Jereb & Roper, 2010) (Moreno et al 2007; Arkhipin 1995; Rocha and Guerra 1999; Arkhipkin 1995; Jereb & Roper, 2010; Natsukari and Komine 1992; Boyle, 1983; Arkhipkin et al., 2015; Jereb et al., 2015; Pierce et al., 2010) 480 “The relationship between age and sexual maturity of male and females in both species [L. forbesii and L. vulgaris] was weak (Fig. 3). Immature and maturing specimens were present at a wide range of ages in both sexes and species…Males and females of L. vulgaris became mature from 300 and 320 days, respectively, and as shown in Fig. 3,they were always found to be mature from the 480 age class (days).” (Agus et al. 2018) (Moreno et al., 2005; Boyle, 1983; Agus et al. 2018; Jereb et al., 2015; Pierce et al., 2010; Arkhipin 1995) 250 "Age studies confirm that males mature ca. 1 month earlier than females (Rocha, 1994;
Arkhipkin, 1995; Bettencourt et al., 1996; Moreno et al., 2005). In Portugal, males mature
at a mean age of 9 months, and spawning takes place at a mean age of 10 months. A
high percentage of the population is mature before 1 year (Moreno et al., 2005). Farther south, on the Saharan Bank, minimum age at full maturity is 250 d in males (ca. 8
months) and 285 d in females (ca. 9.5 months) (Arkhipkin, 1995). " (Jereb et al., 2015)		 (Moreno et al., 2005; Boyle, 1983; Agus et al. 2018; Jereb et al., 2015; Pierce et al., 2010; Arkhipin 1995) 6 5 1 L. vulgaris could feed either in the pelagic zone, or forage along the bottom. This behaviour was supported by the presence of semipelagic species (T. trachurus), pelagic species (S. pilchardus and AtheriNAsp), and benthic species, respectively." (Coelho et al 1997) Coelho et al 1997 1 All the squid sampled used stalking as the hunting mode and approached the prey slowly prior to a sudden strike (Carreno Castilla et al 2020) Carreno Castilla et al 2020 1 squid seized the prey by throwing the tentacles followed by an arm ambush (stalking using a mixed strategy using opening-arms and tentacles (Carreno Castilla 2020) Carreno Castilla 2020
Flores et al., 1983		 1 "Furthermore, by rapidly killing and immobilising the prey, L. vulgaris is able to manipulate and eat the fsh safely, orientating it in such manner, prior to consumption, avoiding potential injuries by the fsh fn spines (Sykes and Gestal 2014)" (Carreno Castilla et al 2020) Sykes and Gestal 2014 Carreno Castilla et al 2020
Iglesias et al., 2014		 1 "Hatchlings of both L. vulgaris and D. opalescens perform external predigestion and ingest only the flesh of their crustacean prey (Boletzky 1974b; Franco-Santos and Vidal in press)" (Iglesias et al., 2014). Iglesias et al., 2014 1 However, distinct diferences in the use of the predation tools for seizing the prey were observed (Carreno Castilla et al 2020)
However, in addition, a fourth phase (iv) post-seizure was identifed. Body patterning performed for each prey type is described for each predation phase (Carrino Castella 2020)		 Carreno Castilla et al 2020
Cabanellos-Reboredo et al 2011		 5 5 1 "squid skin where nerve-evoked colour changes, used in animal-to- animal signalling and camouflage" (Lima et al., 2003) Lima et al., 2003; Hanlon, 1988; Cornwell et al., 1997; Hanlon et al., 1999
Mathger and Denton, 2001		 1 "One of us (L.M.M.) has observed that under stress (e.g. during capture of a squid) the mantle ‘red’ stripes become very prominent, suggesting that iridophores could be controlled by the animal. " (Mathger and Denton, 2001) Mathger and Denton, 2001 1 "The iridophores of the ventral side have high reflectivity in the blue-green at oblique angles of incidence. This will channel the light, which passes through the mantle muscle, downwards, so that the squid minimises the shadow cast below the animal." (Mathger and Denton, 2001) Mathger and Denton, 2001 1 "The squid used in our experiments all appeared very transparent, with the chromatophores mostly in a retracted state." (Mathger and Denton, 2001) Mathger and Denton, 2001 1 “escape reactions, probably including ink-rejection” (Messenger, 1979) Messenger, 1979 1 "Apart from the eyes and the inksac, the body of a squid is very transparent." (Mathger and Denton, 2001) (Mathger and Denton, 2001; Sen, 2006) 37 "Loligo forbesii feeds on small fishes and to a minor extent on other cephalopod species, crustaceans and polychaetes; cannibalism also occurs, but it seems limited to large squids feeding on much smaller ones. Primary prey items vary among geographical areas and seasons. The main food sources are as follows: in Scottish waters whiting (Merlangius merlangius), poor-cod (Trisopterus spp.) and sandeels (Ammodytidae) comprise the most abundant fish species in the diet, whereas in Irish waters sprat (Sprattus sprattus) and poor-cod Trisopterus spp., and off the Azores blue jack mackerel (Trachurus picturatus) dominate. Ontogenetic shifts in feeding habits occur. Crustaceans dominate in the diet of juveniles, but no sex-related nor maturation-related differences occur. " (Jereb & Roper, 2010) Jereb & Roper 2010
Valls et al 2015
Sole et al 2013
Coelho et al 1997
(Iglesias et al., 2014).		 0 0 5 40 Known predators of Loligo vulgaris in the Mediterranean Sea and Northeast Atlantic: Common octopus (Octopus vul-
garis) , Blainville's dogfish (Squalus blainville) , Blackspotted smooth-hound (Mus-
telus punctulatus) , Blonde ray (Raja brachyura) , Bull ray (Pteromylaeus bovinus) , Eagle ray (Myliobatis aquila) , Lesser spotted dogfish (Scyliorhinus
canicula) , Marbled electric ray (Torpedo mar-
morata) , Pelagic stingray (Pteroplatytrygon vi-
olacea) , Smooth-hound (Mustelus mustelus) , Thornback ray (Raja clavata) , Torpedo ray (Torpedo spp.) , Atlantic bluefin tuNA(Thunnus
thynnus) , Atlantic stargazer (Uranoscopus sca-
ber) , Common two-banded seabream
(Diplodus vulgaris) , Greater amberjack (Seriola dumerili) , Lesser weever (Echiichthys vipera) , Spotted flounder (Citharus linguat-
ula) , Swordfish (Xiphias gladius) , Mediterranean monk seal (Mona-
chus monachus) , Harbour porpoise (PhocoeNApho-
coena) , Bottlenose dolphin (Tursiops trunca-
tus) , Common dolphin (Delphinus del-
phis) , Long-finned pilot whale (Globiceph-
alus melas) , Risso's dolphin (Grampus griseus) (Jereb et al., 2015)		 (Blanco et al., 2006) (Jereb
& Roper, 2010) (Arkhipkin
et al., 2015) (Bello
1997) (Di
Lorenzo et al., 2020) (Guclusoy
2008) (Miliou
et al 2007) (Miliou
et al 2006) (Pereira
et al., 1997) (Miokovik
et al 1999) (Matallanas
et al 1995) (Pierce
et al., 2010) (Jereb
et al., 2015)		 5 3 "These squid are schooling animals that need to signal changes in relative position to neighbouring squid to maintain the coherence of the school." (Mathger and Denton, 2001) Mathger and Denton, 2001 1 "These squid are schooling animals that need to signal changes in relative position to neighbouring squid to maintain the coherence of the school." (Mathger and Denton, 2001) Mathger and Denton, 2001 5 0 5 "The fluorescent layers and the iridophores of the ‘red’, ‘green’ and ‘blue’ stripes are orientated in such a way that they reflect light, which always arises from directions in which the radiances are higher than those of the background. They consequently disrupt, rather than aid, camouflage and it seems that their function lies in communication between members of the same species, e.g. signalling between neighbours in schools." (Mathger and Denton, 2001) 1 "In captivity, L. vulgaris mating behaviour and male–male aggression may be induced
by the presence of a recently laid egg mass (or visually similar object) in the tank
or even in the visual field (Arnold 1990), as well as by pheromones present in the
egg mass (King et al. 2003; Cummins et al. 2011). Arnold (1990) observed that on
detecting such an object, individual sexually mature squid investigate the object
tactilely and may jet water at it (possibly an effort to flush away sperm from other
males, see ‘Cuttlefish’ above). Males begin to dart about, display to other males,
and place themselves between females and rival males. Females and males display
to each other, accentuating oviducal gland and testes, respectively ( L. reynaudii,
Hanlon et al. 2002). A male will swim alongside a female and raise one or two me-
dial arm in an S-shaped curved display posture. Dark bands or patches also feature
in this display, especially in competition with rival males, who may also be chased
(Byrne et al. 2003; Mather 2004). Male–male contests may also include physical
contact such as fin beating (Hanlon et al. 2002). Social hierarchies determined via
dominance in agnostic displays develop in captivity (Arnold 1990), while paired
males have an advantage over intruder males in the wild (Hanlon et al. 2002), an
effect due to female choice, as females jet to avoid unwanted male mating attempts.
Copulation of paired L. vulgaris, described by Arnold (1990) and of L. reynau-
dii described by Hanlon et al. (2002), is preceded by the male positioning himself
alongside but slightly below the female, flashing chromatophores. The male then
grabs the female and positions his arms close to her mantle opening. He reaches into
his mantle with his hectocotylus and picks up spermatophores, which are quickly
ejaculated and cemented to the inside of the female mantle near the opening of the
oviduct. The male then releases the female. Copulatory behaviour may be inter-
spersed with egg laying, and newly released sperm may be observed on just-laid
egg masses. Female choice may operate on several levels, including female ma-
nipulation of sex ratios, avoidance of mating attempts, and selection of stored sperm
to fertilize eggs (Hanlon et al. 2002), leading to multiple paternity within egg strings
(Shaw and Sauer 2004). Copulation alternated with egg laying will continue until
both sexes are exhausted. Squid die within hours. Sneaker males of L. reynaudii
(Hanlon et al. 2002), as well as other squid (e.g. D. gigas: Nigmatullin et al. 2001)
and cuttlefish (Hanlon et al. 1999a) mate in the head-to-head position. Mating be-
haviour may be variable and dependent on context (Hanlon et al. 2002; Jantzen and
Havenhand 2003a). Thus, there are opportunities to manipulate the onset of spawn-
ing under cultivation, although courtship and mating behaviours may be inseparable
from male–male aggression" (Iglesias et al., 2014).		 (Iglesias et al., 2014). 1 "In captivity, L. vulgaris mating behaviour and male–male aggression may be induced
by the presence of a recently laid egg mass (or visually similar object) in the tank
or even in the visual field (Arnold 1990), as well as by pheromones present in the
egg mass (King et al. 2003; Cummins et al. 2011). Arnold (1990) observed that on
detecting such an object, individual sexually mature squid investigate the object
tactilely and may jet water at it (possibly an effort to flush away sperm from other
males, see ‘Cuttlefish’ above). Males begin to dart about, display to other males,
and place themselves between females and rival males. Females and males display
to each other, accentuating oviducal gland and testes, respectively ( L. reynaudii,
Hanlon et al. 2002). A male will swim alongside a female and raise one or two me-
dial arm in an S-shaped curved display posture. Dark bands or patches also feature
in this display, especially in competition with rival males, who may also be chased
(Byrne et al. 2003; Mather 2004). Male–male contests may also include physical
contact such as fin beating (Hanlon et al. 2002). Social hierarchies determined via
dominance in agnostic displays develop in captivity (Arnold 1990), while paired
males have an advantage over intruder males in the wild (Hanlon et al. 2002), an
effect due to female choice, as females jet to avoid unwanted male mating attempts.
Copulation of paired L. vulgaris, described by Arnold (1990) and of L. reynau-
dii described by Hanlon et al. (2002), is preceded by the male positioning himself
alongside but slightly below the female, flashing chromatophores. The male then
grabs the female and positions his arms close to her mantle opening. He reaches into
his mantle with his hectocotylus and picks up spermatophores, which are quickly
ejaculated and cemented to the inside of the female mantle near the opening of the
oviduct. The male then releases the female. Copulatory behaviour may be inter-
spersed with egg laying, and newly released sperm may be observed on just-laid
egg masses. Female choice may operate on several levels, including female ma-
nipulation of sex ratios, avoidance of mating attempts, and selection of stored sperm
to fertilize eggs (Hanlon et al. 2002), leading to multiple paternity within egg strings
(Shaw and Sauer 2004). Copulation alternated with egg laying will continue until
both sexes are exhausted. Squid die within hours. Sneaker males of L. reynaudii
(Hanlon et al. 2002), as well as other squid (e.g. D. gigas: Nigmatullin et al. 2001)
and cuttlefish (Hanlon et al. 1999a) mate in the head-to-head position. Mating be-
haviour may be variable and dependent on context (Hanlon et al. 2002; Jantzen and
Havenhand 2003a). Thus, there are opportunities to manipulate the onset of spawn-
ing under cultivation, although courtship and mating behaviours may be inseparable
from male–male aggression" (Iglesias et al., 2014).		 (Iglesias et al., 2014). 1 "In captivity, L. vulgaris mating behaviour and male–male aggression may be induced
by the presence of a recently laid egg mass (or visually similar object) in the tank
or even in the visual field (Arnold 1990), as well as by pheromones present in the
egg mass (King et al. 2003; Cummins et al. 2011). Arnold (1990) observed that on
detecting such an object, individual sexually mature squid investigate the object
tactilely and may jet water at it (possibly an effort to flush away sperm from other
males, see ‘Cuttlefish’ above). Males begin to dart about, display to other males,
and place themselves between females and rival males. Females and males display
to each other, accentuating oviducal gland and testes, respectively ( L. reynaudii,
Hanlon et al. 2002). A male will swim alongside a female and raise one or two me-
dial arm in an S-shaped curved display posture. Dark bands or patches also feature
in this display, especially in competition with rival males, who may also be chased
(Byrne et al. 2003; Mather 2004). Male–male contests may also include physical
contact such as fin beating (Hanlon et al. 2002). Social hierarchies determined via
dominance in agnostic displays develop in captivity (Arnold 1990), while paired
males have an advantage over intruder males in the wild (Hanlon et al. 2002), an
effect due to female choice, as females jet to avoid unwanted male mating attempts.
Copulation of paired L. vulgaris, described by Arnold (1990) and of L. reynau-
dii described by Hanlon et al. (2002), is preceded by the male positioning himself
alongside but slightly below the female, flashing chromatophores. The male then
grabs the female and positions his arms close to her mantle opening. He reaches into
his mantle with his hectocotylus and picks up spermatophores, which are quickly
ejaculated and cemented to the inside of the female mantle near the opening of the
oviduct. The male then releases the female. Copulatory behaviour may be inter-
spersed with egg laying, and newly released sperm may be observed on just-laid
egg masses. Female choice may operate on several levels, including female ma-
nipulation of sex ratios, avoidance of mating attempts, and selection of stored sperm
to fertilize eggs (Hanlon et al. 2002), leading to multiple paternity within egg strings
(Shaw and Sauer 2004). Copulation alternated with egg laying will continue until
both sexes are exhausted. Squid die within hours. Sneaker males of L. reynaudii
(Hanlon et al. 2002), as well as other squid (e.g. D. gigas: Nigmatullin et al. 2001)
and cuttlefish (Hanlon et al. 1999a) mate in the head-to-head position. Mating be-
haviour may be variable and dependent on context (Hanlon et al. 2002; Jantzen and
Havenhand 2003a). Thus, there are opportunities to manipulate the onset of spawn-
ing under cultivation, although courtship and mating behaviours may be inseparable
from male–male aggression" (Iglesias et al., 2014).		 (Iglesias et al., 2014). 1 "In captivity, L. vulgaris mating behaviour and male–male aggression may be induced
by the presence of a recently laid egg mass (or visually similar object) in the tank
or even in the visual field (Arnold 1990), as well as by pheromones present in the
egg mass (King et al. 2003; Cummins et al. 2011). Arnold (1990) observed that on
detecting such an object, individual sexually mature squid investigate the object
tactilely and may jet water at it (possibly an effort to flush away sperm from other
males, see ‘Cuttlefish’ above). Males begin to dart about, display to other males,
and place themselves between females and rival males. Females and males display
to each other, accentuating oviducal gland and testes, respectively ( L. reynaudii,
Hanlon et al. 2002). A male will swim alongside a female and raise one or two me-
dial arm in an S-shaped curved display posture. Dark bands or patches also feature
in this display, especially in competition with rival males, who may also be chased
(Byrne et al. 2003; Mather 2004). Male–male contests may also include physical
contact such as fin beating (Hanlon et al. 2002). Social hierarchies determined via
dominance in agnostic displays develop in captivity (Arnold 1990), while paired
males have an advantage over intruder males in the wild (Hanlon et al. 2002), an
effect due to female choice, as females jet to avoid unwanted male mating attempts.
Copulation of paired L. vulgaris, described by Arnold (1990) and of L. reynau-
dii described by Hanlon et al. (2002), is preceded by the male positioning himself
alongside but slightly below the female, flashing chromatophores. The male then
grabs the female and positions his arms close to her mantle opening. He reaches into
his mantle with his hectocotylus and picks up spermatophores, which are quickly
ejaculated and cemented to the inside of the female mantle near the opening of the
oviduct. The male then releases the female. Copulatory behaviour may be inter-
spersed with egg laying, and newly released sperm may be observed on just-laid
egg masses. Female choice may operate on several levels, including female ma-
nipulation of sex ratios, avoidance of mating attempts, and selection of stored sperm
to fertilize eggs (Hanlon et al. 2002), leading to multiple paternity within egg strings
(Shaw and Sauer 2004). Copulation alternated with egg laying will continue until
both sexes are exhausted. Squid die within hours. Sneaker males of L. reynaudii
(Hanlon et al. 2002), as well as other squid (e.g. D. gigas: Nigmatullin et al. 2001)
and cuttlefish (Hanlon et al. 1999a) mate in the head-to-head position. Mating be-
haviour may be variable and dependent on context (Hanlon et al. 2002; Jantzen and
Havenhand 2003a). Thus, there are opportunities to manipulate the onset of spawn-
ing under cultivation, although courtship and mating behaviours may be inseparable
from male–male aggression" (Iglesias et al., 2014).		 (Iglesias et al., 2014). 0 "without maternal care" (Villanueva & Norman, 2008) Villanueva & Norman, 2008 0 "without maternal care" (Villanueva & Norman, 2008) Villanueva & Norman, 2008 518 413 703.984 447.488 245 166 W-W
Lolliguncula brevis Lolliguncula brevis 26.5 " depths of 8.9 and 26.5 m" (Coehlo et al 2010) (Haimovici et al., 1989; Coehlo et al 2010; Bartol et al 2002; Hanlon et al 1983; Hanlon et al 1983; Vecchione & Roper, 1991, p. 436) 0 0-20 m (Haimovici et al., 1989) (Haimovici et al., 1989; Coehlo et al 2010; Bartol et al 2002; Hanlon et al 1983; Hanlon et al 1983; Vecchione & Roper, 1991, p. 436) 2 "Neritic marine environment, shallw waters from 2 to 8m depth. The sediment varies from sandy to muddy bottoms" (Simone 1997) Jereb & Roper, 2010;
Simone 1997		 73 45 "about 45ºN to 28ºS" (Jereb & Roper, 2010) (Jereb & Roper, 2010) -28 "about 45ºN to 28ºS" (Jereb & Roper, 2010) (Jereb & Roper, 2010) 1 1 "Typically neritic, in the northwest Atlantic, North of Cape Hatteras, D. pealeii migrates inshore and northward in late spring and early summer into shallow coastal waters to spawn; by late autumn to early winter the squid migrate southward and into deeper waters along the edge of the continental shelf, where they over-winter. These inshore-offshore and north-south movements are mainly related to the avoidance of water temperatures of 8°C or below. Large concentrations of squid are associated with frontal zones with strong temperature gradients, and they are concentrated mostly along the warm-water side. Hence the definition of “member of the migratory, warm-water group of species, centred primarily in mid-Atlantic waters” (Murawski, 1993), which makes seasonal migrations." (Jereb & Roper, 2010) (Jereb & Roper, 2010; Arkhipkin et al., 2015; Staudinger et al. 2006; Granados-Amores et al 2019; Lange & Sissenwine 1980; Boyle, 1983) 1 "Typically neritic, in the northwest Atlantic, North of Cape Hatteras, D. pealeii migrates inshore and northward in late spring and early summer into shallow coastal waters to spawn; by late autumn to early winter the squid migrate southward and into deeper waters along the edge of the continental shelf, where they over-winter. These inshore-offshore and north-south movements are mainly related to the avoidance of water temperatures of 8°C or below. Large concentrations of squid are associated with frontal zones with strong temperature gradients, and they are concentrated mostly along the warm-water side. Hence the definition of “member of the migratory, warm-water group of species, centred primarily in mid-Atlantic waters” (Murawski, 1993), which makes seasonal migrations." (Jereb & Roper, 2010) (Jereb & Roper, 2010) 365.3 "L. brevis displays a shorter life expectancy of ~1 year
(Hixon et al., 1981)" (Zielinski and Poertner 2000)		 (Jackson et al 1997; Zielinski and Poertner 2000; Pattillo et al 1997; Hanlon et al 1983) 48 "Males and females survived equally
well during these observations; overall mean survival was 50 days (Sx = 4.4) for males
and 48 days (Sx = 4.1) for females" (Hanlon et al 1983)		 (Jackson et al 1997; Zielinski and Poertner 2000; Pattillo et al 1997; Hanlon et al 1983) 200 Sexual maturity: "in less than 200 days" (Perez & Zaleski, 2013) (Perez & Zaleski, 2013) 200 Sexual maturity: "in less than 200 days" (Perez & Zaleski, 2013) (Perez & Zaleski, 2013) 5 5 1 here they often display a very unusual behaviour, mimicking the colour, form and swimming pattern of the small fishes nearby, probably as an effective form of hunting technique as previously observed in another squid, Sepioteuthis sepioidea. (Jereb & Roper, 2010). Jereb & Roper, 2010
(Hanlon & Messenger, 1996)." (Martins & Perez, 2006, p 32		 1 "fin braking is integral for turning maneuvers, with kinematic studies showing that fin activity in L. brevis is important for tight and rapid turns (Jastrebsky et al., 2016) and for pursuing and capturing prey (Jastrebsky et al., 2017)" (Bartol et al 2018) Bartol et al 2018 1 Because prey strikes occur in an arms-first orientation (Hanlon and Messenger, 1996; Kier and van Leeuwen, 1997), arms-first swimming enables swift prey attacks whenever opportunities arise, while tail-first swimming requires rotation to arms-first swimming prior to attack." (Bartol et al 2001) Bartol et al 2001
Pattillo et al 1997		 1 Prey items (e.g. fish) are injected with venom usually through bites behind the head with the squid's parrot-like beak. The venom acts as a tranquilizer that paralyzes the prey. (Pattillo et al 1997) Pattillo et al 1997 1 Once fish prey are paralyzed, the squid consumes the viscera, and then strips the flesh from the animal by means of perforating bites down the animal's sides" Pattillo et al 1997 Pattillo et al 1997 0 Squid attack success was high (>80%) and three behavioral phases were identified: (1) approach, (2) strike and (3) recoil. […] . Irrespective of prey type, L. brevis consistently positioned themselves above the prey target prior to the tentacle strike, possibly to facilitate a more advantageous downward projection of the tentacles" […] "agility, maneuverability, and swimming speed/acceleration are all important for prey capture in L. brevis" […] " squid located prey positioned laterally to their bodies, turned and attacked head first (and eyes first). (Jastrebsky et al 2017) Jastrebsky et al 2017 6 6 1 " Furthermore, it was suggested that it dives into hypoxic
waters to avoid predators and/or to feed (Vecchione 1991a)" Zielinski et al 2000 		Pörtner et al. 2002
Mederios Moraes and Lavrado 2017
Vecchione 1991a
Zielinski et al 2000		 1 "that clear body patterns were used significantly more often than intermediate body patterns and dark body patterns . The velocity of the approaching predator did not affect the body pattern selection of the paralarvae (all p N 0.05), nor did the angle of the predator approach, distance of the predator or distance traveled by the predator at the time of the interaction (all p N 0.05). When responding to an approaching predator, juvenile and adult squid were significantly more likely to demonstrate the banded pattern than the dark body, dark arms with clear body, or clear body pattern (ANOVA: F3,76 = 26.1, p b 0.001; Fig. 9B). When the juvenile/adult squid responded with an inking event (something that is very rare for paralarvae), it was significantly more likely to be in the form of a ‘pseudomorph’ than a ‘cloud’ or ‘puff‘(ANOVA: F2,57 = 91.2, p b 0.001)" (York and Bartol 2016) York and Bartol 2016 1 "The squid used in our experiments all appeared very transparent, with the chromatophores mostly in a retracted state." (Mathger and Denton, 2001) Mathger and Denton, 2001 1 The function of this body posture is thought to be camouflage, allowing the squid to thwart predators by mimicking floating debris, such as drifting seaweed pieces (Hanlon & Messenger, 1996). Hanlon & Messenger, 1996 1 Lolliguncula brevis (Dubas et al., Reference Dubas, Hanlon, Ferguson and Pinsker1986) and L. panamensis (Moynihan & Rodaniche, Reference Moynihan and Rodaniche1982) tend to show displays intermediate between the Deimatic and the Flamboyant (described below); that is, they have arms flared in Upward V-curl, light bodies with some sort of dark border (on the fins or the flared arms), but they do not have conspicuous false eyespots." (Hanlon & Messenger, 2018, p. 125) Hanlon & Messenger, 2018, p. 125
Hanlon &
Messenger, 1996		 1 "The juvenile and adult squid in this study responded to an oncoming predator with an inking event in approximately 60% of all interactions. Inking events were always exhibited in sequence with an escape jet, where inking occurred either at the initiation of the escape jet (61%) or at another point throughout the escape jet (39%). Inking in paralarvae only occurred in 2.0% of all interactions." (York and Bartol 2016) York and Bartol 2016 7 "Food consists of small crustaceans and fishes" (Jereb & Roper, 2010) Jereb and Roper 2010
Bartol et al 2018
Coehlo et al 2010
Pattillo et al 2017		 0 1 Coehlo et al 2010 4 8 "GuiaNAdolphins Sotalia guianensis and franciscanas Pontoporia blainvillei" (Cremer et al 2012) (Jereb
& Roper, 2010) (Bornatowski
et al 2012) (Mederios
Moraes and Lavrado 2017) (Cremer
et al 2012) (York
and Bartol 2014) (Pate
and McFee 2012) (Turner
and Rooker 2005) (Lessa
and Almeida 1997) 		4 3 "Lolliguncula brevis is a social, schooling species that spends most of its time in shallow water near the bottom" Dubas et al 1986 Dubas et al 1986 1 "Lolliguncula brevis is a social, schooling species that spends most of its time in shallow water near the bottom" Dubas et al 1986 Dubas et al 1986 6 5 1 "Habituation and dishabituation are demonstrated for the first time in a squid (Cephalopoda:
Teuthoidea). Each squid (n = 29) was exposed briefly to a plastic model of a predator to determine
how the behavioural responses changed with repeated trials at 1 min intervals. Behavioural responses
were video-monitored for subsequent measurements of the number of escape jets and the duration of
neurally controlled body pattern rings on each trial. Squids habituated readily to a teleost fish model
and could differentiate clearly between a teleost fish and a shark model of the same size. Following a
single series of 15 trials, habituated responses recovered after a 1 hr rest. Habituated responses also
recovered rapidly (dishabituation) when a noxious stimulus was presented. Spaced Training was more
effective than Massed Training in producing longer-term habituation." Long et al 1989		 Long et al 1989 1 Habituation; Classical conditioning (Marini et al. 2017, Table 3) Marini et al. 2017, Table 3 1 "Habituation and dishabituation are demonstrated for the first time in a squid (Cephalopoda:
Teuthoidea). Each squid (n = 29) was exposed briefly to a plastic model of a predator to determine
how the behavioural responses changed with repeated trials at 1 min intervals. Behavioural responses
were video-monitored for subsequent measurements of the number of escape jets and the duration of
neurally controlled body pattern rings on each trial. Squids habituated readily to a teleost fish model
and could differentiate clearly between a teleost fish and a shark model of the same size. Following a
single series of 15 trials, habituated responses recovered after a 1 hr rest. Habituated responses also
recovered rapidly (dishabituation) when a noxious stimulus was presented. Spaced Training was more
effective than Massed Training in producing longer-term habituation…Lolliguncula brevis distinguished the fish models easily although they were identical in size, but thiswas not surprising in view of their keen visual acuity and discrimination (Allen et al, 1985; Flores, 1983; Messenger, 1981)" Long et al 1989		 Long et al 1989 1 "Short-Term HabituationTwenty animals were run through 15 trials of presenting a model fish predator at lmin interstimulus intervals (ISI) and they habituated quickly (Figure 2).Although there was variability among the animals, repeated presentation of themodel predator brought about gradually diminishing responses in both jets and rings" Long et al 1989 Long et al 1989 1 "The longer-term habituation study emphasized the efficacy of space training. It is unlikely that a squid in the wild will be continuously exposed to a stimulus with equal ISIs; stimuli will be irregularly spaced. Spaced training was far more effective in producing longer-term memory than massed training (Figure 5). The superiority of spaced training in producing longer-term habituation is similar to the learning process in vertebrates (Kandel, 1976)" Long et al 1989 Long et al 1989 1 1 "The results of an interspecific association analysis (Cox, 1980 based on Cole, 1949)
based on 150 trawl stations showed that there was a positive coefficient of association
between Lolliguncula brevis and Loligo plei, indicating that these species are found
frequently in close proximity to one another." (Hanlon et al 1983)		 Hanlon et al 1983 0 "Little intraspecific aggression has been
observed and there has been no evidence of rank ordering among males" (Hanlon et al 1983) & " Lolliguncula brevis did not show obvious signs of
(intraspecific) aggression" (Hanlon et al 1983)		 (Hanlon et al 1983) 115 110 30.8 75 NY
Lycoteuthis lorigera Lycoteuthis lorigera 1388 "This species was present in trawls between 192 and 1388 m deep, but was absent on the continental shelf." (Hoving et al., 2007) (Hoving et al 2014; Forch and Uozomi 1990; Cairns 1976; Hoving et al., 2007) 366 "Adults have been captured with open nets between 366-589 m and juveniles have been taken between 46-57 m (Voss, 1962)." (Cairns 1976) (Hoving et al 2014; Forch and Uozomi 1990; Cairns 1976; Hoving et al., 2007) 1 "Oceanic" (Gomes-Pereira et al., 2016) Gomes-Pereira et al., 2016 40 -15 15 S to 55 S (Jereb & Roper, 2010) (Jereb & Roper, 2010) -55 15 S to 55 S (Jereb & Roper, 2010) (Jereb & Roper, 2010) 2 386 "Only one mature male (ML 160 mm) was aged, and preliminary estimates suggest it was 386 days old." (Hoving et al., 2007) (Hoving et al 2007) 131 "Estimates of age ranged from 131 to 315 days and varied with mantle length." (Hoving et al., 2007) (Hoving et al 2007) 334.8 "On the South Atlantic continental slope of South Africa, female Lycoteuthis lorigera reach maturity around 11 months (ML 100 mm) and have been estimated to live for 1 year (Hoving et al., 2007)"(Elsevier, 2014). (Hoving et al., 2014; Hoving et al., 2007; Elsevier, 2014) 299 "The same was done to estimate the age at maturity for females (=299 days)." (Hoving et al., 2007) (Hoving et al., 2014; Hoving et al., 2007; Elsevier, 2014) NA 0 NA 0 3 "This species feeds on pelagic crustaceans and fishes, including myctophids (Voss 1962)" (Hoving et al 2007) Hoving et al 2017
Voss (1962) and Lipinski (1992)		 0 0 2 12 "L. lorigera is preyed upon by commercially
important fish, such as deep-water Cape hake,
Merluccius paradoxus, and Kingklip, Genypterus
capensis (Lipinski et al. 1992), as well as several
smaller cetacean species (Ross 1984), the Portuguese shark Centroscymnus coelolepis (Ebert
et al. 1992), ribbonfish Lepidopus caudatus
(Meyer and Smale 1991), the southern lanternshark Etmopterus granulosus (Lipinski et al.
1992) and in New Zealand waters the species is
preyed upon by petrels (Imber 1975)." (Hoving et al 2007)		 (Young
& Cockcroft 1995) (van den Hoff 2004) (Petry et al 2008) (Hoving et al
2007) (Hoving et al., 2014)		 4 0 not gregarious (inferred from photo and video material) NA 0 NA 21 10 189.8 80 NY
Macrotritopus defilippi Macrotritopus defilippi 1500 "Only 5 of 17 Macrotritopus defilippi paralarvae (4–12 mm ML)
were collected in day tows at the surface (0–200 m) while during the night tows, 12 specimens were collected from 0
to 1500 m." (Judkins & Vecchione, 2020)		 (Judkins & Vecchione, 2020; Haimovici et al., 1989; Jereb et al., 2014; Gerovasileiou et al 2020; Crocetta et al 2015; Guerra et al 2013; Soriano et al 2003; Vecchione & Roper, 1991, p. 438; El-Ganainy and Riad 2008) 0 "Only 5 of 17 Macrotritopus defilippi paralarvae (4–12 mm ML)
were collected in day tows at the surface (0–200 m) while during the night tows, 12 specimens were collected from 0
to 1500 m." (Judkins & Vecchione, 2020)		 (Judkins & Vecchione, 2020; Haimovici et al., 1989; Jereb et al., 2014; Gerovasileiou et al 2020; Crocetta et al 2015; Guerra et al 2013; Soriano et al 2003; Vecchione & Roper, 1991, p. 438; El-Ganainy and Riad 2008) 2 benthic (Chesalin & Zuyev, 2002) Chesalin & Zuyev, 2002
Haimovici et al., 1989
Guerra et al 2013		 38 68 68 N to 30 N (Jereb et al., 2014) (Jereb et al., 2014) 30 68 N to 30 N (Jereb et al., 2014) (Jereb et al., 2014) 1 "Additional material needs to be collected to make firm
conclusions on vertical migration and any possible ontogenic
shift " (Judkins & Vecchione, 2020)		 (Judkins & Vecchione, 2020) 1095.8 "The evolution of the population structure examined suggests a life-span of
about 2 years and a maximum age of about 3 years for combined sexes." (El-Ganainy & Raid, 2008)		 (El-Ganainy & Raid, 2008) 730.5 "The evolution of the population structure examined suggests a life-span of
about 2 years and a maximum age of about 3 years for combined sexes." (El-Ganainy & Raid, 2008)		 (El-Ganainy & Raid, 2008) 4 4 1 The two most frequently used behaviors during foraging events for both species were speculative bottom searching and sitting. (Bennice et al 2021) Bennice et al 2021 1 M. defilippi were flounder mimicry swimming, tripod stance, and grope searching. (Bennice et al 2021) Bennice et al 2021 1 M. defilippi were flounder mimicry swimming, tripod stance, and grope searching. (Bennice et al 2021) Bennice et al 2021 1 "The two most frequently used behaviors during foraging events for both species were speculative bottom searching and sitting." (Bennice et al 2021) Bennice et al 2021 6 5 1 "M. defilippi [hides] in holes and burrows in the sand" (Bennice et al., 2021) Bennice et al., 2021
Hanlon et al 2013
Hanlon & Messenger, 2018
Hochberg & Couch; Hanlon, Reference Hanlon1988; Hanlon, Watson & Barbosa, Reference Hanlon, Watson and Barbosa2010		 1 well camouflaged when stationary, and details of camouflaging techniques are described for M. defilippi. Octopuses implemented flounder mimicry only during swimming, when their movement would give away camouflage in this open sandy habitat. Thus, both camouflage and fish mimicry
were used by the octopuses as a primary defense against
visual predators" (Hanlon et al 2010)		 Hanlon et al 2010
Hanlon et al 2013		 1 "O. vulgaris in the Caribbean sometimes modifies its body posture and skin papillae to conduct a ‘moving algae’ walking behaviour when the surrounding objects on a sand plain are spiky forms of algae and soft corals (Fig. 5.12). This latter behaviour has also been observed – in modified form – in Macrotritopus defilippi in Florida, O. rubescens in California (while swimming), and Abdopus aculeatus in Australia (Fig. 5.11)." (Hanlon & Messenger, 2018, p. 111) Hanlon & Messenger, 2018 1 "M. defilippi used flounder swimming to move across soft bottom and mimic a flatfish that inhabits this lagoon to avoid attack by predators. This flatfish species (Bothus lunatus) is not poisonous and likely not a case of Batesian mimicry or Müllerian mimicry; however, it is unknown if M. defilippi is poisonous. A possible explanation of flatfish mimicry is that a smaller gape predator that could bite a portion of a malleable octopus would mistake the octopus for a larger, bony flatfish that would be too large for its mouth (Huffard, 2006; Hanlon et al., 2010)" (Bennice et al., 2021) [Location South Florida lagoon] Huffard, 2006; Hanlon et al., 2010
Bennice et al., 2021
Hanlon et al 2010
Hanlon & Messenger, 2018		 1 (Jereb et al., 2014) 1 "When attacked, these octopuses are capable of autotomising their arms at the base: the writhing severed arm acting as a decoy to predators and aiding escape. Lost arms regenerate within 2–3 months." (Norman & Finn 2001) Norman & Finn 2001 2 "Only one bivalve and zero gastropods were recorded for M. defilippi. The dominant prey from each category was Calappa spp. (crustacean, excluding unidenti- fied crustaceans) and Laevecardium mortoni (bivalve)." (Bennice et al., 2021) [Location South Florida lagoon] Bennice et al., 2021 0 0 2 2 found
in stomachs of bullet tuNAAuxis rochei Tyrrhenian Sea (Mostarda et al. 2007) 		(Mostarda
et al. 2007) (Hanlon et al 2010)		 1 0 not gregarious (inferred from photo and video material) 1 0 1 1 "Amphioctopus spp., Macrotritopus defilippi, H. maculosa and Wonderpus
photogenicus, which all live in sand or silt habitats, carry their eggs
in the ventral aboral web, in line of water expelled from the funnel
(Tranter and Augustine, 1973;Hanlon et al., 1985;Huffard and Hochberg, 2005;Miske and Kirchhauser, 2006)" (Morse & Huffard, 2019)		 (Morse & Huffard, 2019) 0 "Amphioctopus spp., Macrotritopus defilippi, H. maculosa and Wonderpus
photogenicus, which all live in sand or silt habitats, carry their eggs
in the ventral aboral web, in line of water expelled from the funnel
(Tranter and Augustine, 1973;Hanlon et al., 1985;Huffard and Hochberg, 2005;Miske and Kirchhauser, 2006)" (Morse & Huffard, 2019)		 (Morse & Huffard, 2019) 26 14 124.032 76.976 36.4 40 25 W-W-NY
Mastigoteuthis schmidti Mastigoteuthis agassizii 2250 inferred from closely related species (Boyle & Rodhouse, 2005) 500 inferred from closely related species (Boyle & Rodhouse, 2005) 1 inferred from ROV data of closely related whip-lash squids (see e.g., https://www.youtube.com/watch?v=q2XQ61Wudak) 110 60 "Mastigoteuthis schmidti (Fig. 24.22) is widely distributed between 60"“N and 50“S (Clarke 1966, Clarke and Lu 1974, 1975, Lu and Clarke 1975a, 1975b, R.E. Young 1978)." (Nixon & Young 2003) (Nixon & Young 2003) -50 "Mastigoteuthis schmidti (Fig. 24.22) is widely distributed between 60"“N and 50“S (Clarke 1966, Clarke and Lu 1974, 1975, Lu and Clarke 1975a, 1975b, R.E. Young 1978)." (Nixon & Young 2003) (Nixon & Young 2003) 4 1 1 1 "Mastigoteuthis sp., photographed from a submersible, shows the extremely long tentacles emerging from their tentacular sheaths; these are formed by the enrolled lateral membranes of ventral arms lV, with which the tentacles appear to be in continuity. The tentacles, held apart by the ventral arms, are trailing close to, or over, the substrate [R.E. Young er al. 1998b}. This would allow the very small and very numerous suckers to detect and capture small bottom associated zooplankton." (Nixon & Young 2003) Nixon & Young 2003 NA 0 0 0 2 "found
in stomachs of Cuvier’s beaked whales (Ziphius cavirostris)" (Santos et al.
2001)		 (Santos et al. 2001) (Santos
et al. 2007) (Fernandez
et al., 2014)		 1 0 not gregarious (inferred from photo and video material) NA 0 NA 3 3 28.6 60 NY
Megalocranchia maxima Megalocranchia oceanica 2000 For the genus: "Paralarvae are epipelagic from very near-surface waters to about 200 m day and night. Juveniles metamorphose at about 40 to 50 mm mantle length, are concentrated in the upper 200 m at night, then descend to 600 to 800 m during the day. Subadults undergo further ontogenetic vertical descent through the mesopelagic and into the bathypelagic zone to greater than 2 000 m during daytime and some appear to migrate at night back toward lower epipelagic and mesopelagic depths (about 100 to 700 m). While adults probably mature in the bathypelagic water, they apparently reverse migrate back into epipelagic waters to spawn, as indicated by specimens caught at or near the surface at night" (Jereb & Roper, 2010) (Imber 1978; Jereb & Roper, 2010) 100 For the genus: "Paralarvae are epipelagic from very near-surface waters to about 200 m day and night. Juveniles metamorphose at about 40 to 50 mm mantle length, are concentrated in the upper 200 m at night, then descend to 600 to 800 m during the day. Subadults undergo further ontogenetic vertical descent through the mesopelagic and into the bathypelagic zone to greater than 2 000 m during daytime and some appear to migrate at night back toward lower epipelagic and mesopelagic depths (about 100 to 700 m). While adults probably mature in the bathypelagic water, they apparently reverse migrate back into epipelagic waters to spawn, as indicated by specimens caught at or near the surface at night" (Jereb & Roper, 2010) (Imber 1978; Jereb & Roper, 2010) 1 For the genus: "Paralarvae are epipelagic from very near-surface waters to about 200 m day and night. Juveniles metamorphose at about 40 to 50 mm mantle length, are concentrated in the upper 200 m at night, then descend to 600 to 800 m during the day. Subadults undergo further ontogenetic vertical descent through the mesopelagic and into the bathypelagic zone to greater than 2 000 m during daytime and some appear to migrate at night back toward lower epipelagic and mesopelagic depths (about 100 to 700 m). While adults probably mature in the bathypelagic water, they apparently reverse migrate back into epipelagic waters to spawn, as indicated by specimens caught at or near the surface at night" (Jereb & Roper, 2010) Jereb & Roper, 2010 106 53 "The data indicate that this species is cosmopolitan between 53°N (Iwai 1956a) and 53°S (ex royal albatross. Campbell Island). However, it has been taken most abundantly within latitudes 20-38°." (Imber 1978) (Imber 1978) -53 "The data indicate that this species is cosmopolitan between 53°N (Iwai 1956a) and 53°S (ex royal albatross. Campbell Island). However, it has been taken most abundantly within latitudes 20-38°." (Imber 1978) (Imber 1978) 3 1 For the genus: "Paralarvae are epipelagic from very near-surface waters to about 200 m day and night. Juveniles metamorphose at about 40 to 50 mm mantle length, are concentrated in the upper 200 m at night, then descend to 600 to 800 m during the day. Subadults undergo further ontogenetic vertical descent through the mesopelagic and into the bathypelagic zone to greater than 2 000 m during daytime and some appear to migrate at night back toward lower epipelagic and mesopelagic depths (about 100 to 700 m). While adults probably mature in the bathypelagic water, they apparently reverse migrate back into epipelagic waters to spawn, as indicated by specimens caught at or near the surface at night" (Jereb & Roper, 2010) (Jereb & Roper, 2010) 1 For the genus: "Paralarvae are epipelagic from very near-surface waters to about 200 m day and night. Juveniles metamorphose at about 40 to 50 mm mantle length, are concentrated in the upper 200 m at night, then descend to 600 to 800 m during the day. Subadults undergo further ontogenetic vertical descent through the mesopelagic and into the bathypelagic zone to greater than 2 000 m during daytime and some appear to migrate at night back toward lower epipelagic and mesopelagic depths (about 100 to 700 m). While adults probably mature in the bathypelagic water, they apparently reverse migrate back into epipelagic waters to spawn, as indicated by specimens caught at or near the surface at night" (Jereb & Roper, 2010) (Jereb & Roper, 2010) NA 0 NA 0 1 ink sac (unique within the family) (Jereb & Roper, 2010) (Jereb & Roper, 2010; Nesis, 1974) 0 0 4 "It has been recorded frequently from sperm whales' stomachs (lwai 1956a, Okutani et al. 1976, Clarke et al. 1976), and my studies show that it is taken by certain procellariiform seabirds." beaks found in petrels and albatrosses New Zealand waters (Imber 1978) (Imber 1978) (Vaske
Júnior et al. 2009)		 2 0 not gregarious (inferred from photo and video material) NA 0 NA 3 1 198.4 335 NY
Neorossia caroli Neorossia caroli 1744 "Depth range from 40 to 1744 m. Neorossia caroli is the most bathyal among the species belonging to the family Sepiolidae, collected down to the greatest depths in the western Mediterranean basin (1744 m) and in the eastern Atlantic (1535 m)." (Jereb & Roper, 2005) (Jereb & Roper, 2005; Quetglas et al., 2000; Villanueva 1992; Goren et al 2006; Villanueva 1992; Cuccu et al 2007; Salman 2011; Krstulovic Sifner et al 2007) 40 "Depth range from 40 to 1744 m. Neorossia caroli is the most bathyal among the species belonging to the family Sepiolidae, collected down to the greatest depths in the western Mediterranean basin (1744 m) and in the eastern Atlantic (1535 m)." (Jereb & Roper, 2005) (Jereb & Roper, 2005; Quetglas et al., 2000; Villanueva 1992; Goren et al 2006; Villanueva 1992; Cuccu et al 2007; Salman 2011; Krstulovic Sifner et al 2007) 2 "It is a demersal species living preferentially on deep muddy bottoms characterized by Isidella elongata populations, often overlapping with Rossia macrosoma in the upper level of its distributional range and frequently associated with Sepietta oweniaNAand Rondeletiola minor." (Jereb & Roper, 2005) Jereb & Roper, 2005 120 65 65 N to 55 S (Jereb & Roper, 2005) (Jereb & Roper, 2005) -55 65 N to 55 S (Jereb & Roper, 2005) (Jereb & Roper, 2005) 4 1 "The occurrence of small individuals of both N. caroli and B. sponsalis at greater depths than larger individuals, suggests there may be an upslope ontogenetic migration." (Jereb & Roper, 2005) (Jereb & Roper, 2005; Villanueva 1992) 730.5 "The lifespan is probably between 12 and 24 months." (Jereb & Roper, 2005) (Jereb & Roper, 2005) 365.3 "The lifespan is probably between 12 and 24 months." (Jereb & Roper, 2005) (Jereb & Roper, 2005) NA 0 NA 0 0 "the ink sac is reduced and does not contain ink while it lacks also a luminous organ." (Goren et al 2006) (Goren et al 2006; Jereb & Roper, 2005; von Boletzky ,1971) 0 "the ink sac is reduced and does not contain ink while it lacks also a luminous organ." (Goren et al 2006) (Goren et al 2006; Jereb & Roper, 2005; von Boletzky ,1971) 0 0 3 "made
up about 2% of cephalopod species found in stomachs of demersal sharks
Scyliorhinus canicula caught in Aegean Sea" (Kousteni et al. 2018)		 (Kousteni
et al. 2018) (Jereb
et al., 2015) (Bello
1997) 		2 0 not gregarious (inferred from photo and video material) NA 0 NA 19 16 21.4 60 NY
Neoteuthis thielei Neoteuthis thielei 2000 "The ALAMINOS specimens are the first recorded for the Gulf and for the western North Atlantic. The Gulf specimens were collected in open waters; the smaller one (ML 45) at a discrete depth of 900 meters, the larger in a net fished to 2000 meters." Lipka 1975: 146 Lipka 1975 448 448 m (Vecchione & Roper, 1991, p. 436) (Vecchione & Roper, 1991, p. 436) 1 "Oceanic, mesopelagic " (Gomes-Pereira et al., 2016) Gomes-Pereira et al., 2016 125 70 70 N to 55 S (Jereb & Roper, 2010) (Jereb & Roper, 2010) -55 70 N to 55 S (Jereb & Roper, 2010) (Jereb & Roper, 2010) 4 1 "Paralarvae and juveniles epipelagic to mesopelagic; adults are mesopelagic to bathypelagic." (Jereb & Roper, 2010) (Jereb & Roper, 2010) NA 0 NA 0 0 0 0 not gregarious (inferred from photo and video material) NA 0 NA 1 0 18 23 NY
Octopoteuthis danae Octopoteuthis danae 1000 Caught from 0-1000 m (Herring et al., 1992) (Jereb & Roper, 2010; Herring et al., 1992) 0 Caught from 0-1000 m (Herring et al., 1992) (Jereb & Roper, 2010; Herring et al., 1992) 1 inferred from ROV material 36 32 Caught from 32" 32 N to 4" 50 S (Herring et al., 1992) (Herring et al., 1992) -4 Caught from 32" 32 N to 4" 50 S (Herring et al., 1992) (Herring et al., 1992) 4 1 "The young are found in surface waters and the adults inhabit
the deep sea." (Nixon & Young 2003)		 (Nixon & Young 2003) NA 0 NA 0 1 Present in the family (Jereb & Roper, 2010) (Jereb & Roper, 2010) 0 0 1 found in stomachs of blue shark off Ivory Coast (Konan et al. 2018) (Konan et al. 2018) 1 0 not gregarious (inferred from photo and video material) NA 0 NA 1 4 139.6 70 NY
Octopus bimaculatus Octopus bimaculatus 212 “were collected in depths of 208–212 m, much deeper than the recognised depth limit for this species (i.e., 50 m) (Jereb et al. 2014)” (Urbano and Hendrickx, 2018) (Villanueva, 2008; Jereb et al., 2014; Urbano & Hendrickx, 2018) 0 "Depths range from
0 to 50 m." (Jereb et al., 2014)		 (Villanueva, 2008; Jereb et al., 2014; Urbano & Hendrickx, 2018) 2 "While many octopuses live on rocky or coral reefs, others are known to inhabit kelp holdfasts, sand, or mud habitats[…] see Am- brose, 1988)." Ambrose, 1988 16 35 35 N to 20 N (Jereb et al., 2014) (Jereb et al., 2014) 19 "Conception Point (34°26.53′ N), USA to Manzanillo (19°04.09′ N), Mexico" (Urbano & Hendrickx, 2019) (Urbano & Hendrickx, 2019) 1 730.5 Life span: 1-2 years (Ambrose, 1988) (Ambrose 1990 in Dominguez-Contreras et al. 2018; Ambrose, 1988) 365.3 Life span: 1-2 years (Ambrose, 1988) (Ambrose 1990 in Dominguez-Contreras et al. 2018; Ambrose, 1988) 517.4 Sexual maturity: 12-17 months (Ambrose, 1988) (Ambrose, 1988) 365.25 Sexual maturity: 12-17 months (Ambrose, 1988) (Ambrose, 1988) 5 5 1 foraging behavior (Hofmeister 2018) Hofmeister 2018 1 The second octopus repeatedly stalked other scorpionfish. (Taylor & Chen 1969) Taylor & Chen 1969 1 "Speculative hunting
has been described as a pounce (on an object such as kelp, a coral head or a
rock) with outspread web and then a feel around under the web for the presence
of food items (Yarnall, 1969). This type
of hunting has also been reported for […] Octopus bimaculatus (R.F. Ambrose, personal
communication in Hanlon &
Messenger, 1996)." (Cosgrove, 2002)		 Hanlon & Messenger, 1996)
(Cosgrove, 2002)
Hanlon & Messenger, 2018, p. 84		 1 An octopus captures a scorpionfish by attaching two or more arms along the fish's body, hauling it toward the horny beak, and enveloping it with more arms and the web. (Taylor & Chen 1969) Taylor & Chen 1969 1 Drilling is well documented (Hiemstra, 2015) Hiemstra, 2015
Ambrose 1986
Schmitt et al. 1983
Casey, 1999 in Anderson et al., 2008
Pilson & Taylor 1961		 4 3 1 "Previous research has determined that O. bimaculatus individuals can occupy dens for 1 to 5 months (Ambrose, 1982, 1983)." (Hofmeister & Voss, 2017) Hofmeister & Voss, 2017
Hanlon & Messenger, 2018, p. 117-118
Yarnall (1969) in Hofmeister & Voss, 2017		 1 "On a number of occasions octopuses were observed to dig or blow out shelters. This is accomplished by a combination of jets of water directed forcefully at the sand and movement of sand by suckers, arms, and web. Excavated sand is dispersed by forceful jets of water. The process is quite rapid, and a hole up to 20 cm deep may be dug in a matter of minutes. All observed cases of digging involved cracks in bedrock which were filled with sand and shelly debris; shelters are also dug in sand. I have not observed complete shelter construction, but some shelter modification such as moving loose stones or debris seems to be universal." (Ambrose 1982) Ambrose 1982
Alves et al., 2008		 1 Deimatic Displays in other octopuses differ slightly. For example, Octopus maya has diffuse ‘spots’ but no distinct ocelli and Octopus bimaculoides, O. bimaculatus and O. filosus have prominent ocelli at the bases of their arms that are used with lightly graded mottles to produce a Deimatic Display. " (Hanlon & Messenger, 2018, p. 125) Hanlon & Messenger, 2018, p. 125 1 (Jereb et al., 2014; Herring et al., 1992) 59 "O. bimaculatus consumes a wide variety of invertebrate prey items including bivalves, gastropods, crustaceans, small fish, and other octopods, and can affect the abundance and diversity of prey species (Ambrose, 1984, 1986)." (Hofmeister & Voss 2017) Jereb et al 2014
Hofmeister 2018
Hofmeister & Voss 2017
Armendariz Villegas et al. 2014
Ambrose 1984		 0 0 5 4 Santa
Catalina, CA: "Only the kelp bass, Paralabrax clathratus (Girard, 1854),
and the California sheephead, Semicossyphus pulcher (Ayres, 1854) were included
as they are the two primary fish predators of octopuses in this area. California moray eels, Gymnothorax mordax"
(Ayres, 1859) (Hofmeister 2018)		 (Hanlon
& Messenger, 2018, p. 145) (Taylor
& Chen 1969) (Lopez-Peraza
et al. 2017) (Hofmeister
2018) (Villanueva
& Norman, 2008)		 1 2 "The existence of a population comprised mainly of permanent residents has important implications for the ecology of Octopus bimaculatus. Mating and agonistic behavior may be influenced by familiartiy between participants. A non-transient octopus population could result in a different pattern or intensity of resource utilization than a migratory or purely transient population." (Ambrose 1982) Ambrose 1982 0 "The existence of a population comprised mainly of permanent residents has important implications for the ecology of Octopus bimaculatus. Mating and agonistic behavior may be influenced by familiartiy between participants. A non-transient octopus population could result in a different pattern or intensity of resource utilization than a migratory or purely transient population." (Ambrose 1982) Ambrose 1982 10 8 1 (On one specimen of bimaculatus, one of vulgaris) "Both animals were fed with crabs and pieces of shrimp on the subsequent days and by August 11th (=Day 1) were taking them readily and training was begun with smooth plastic black and while balls 12 mm in diameter. The octopuses were rewarded with a piece of shrimp for each attack at either ball, if it was taken under the web. The two colours were given alternately, each for either three or four trials a day. From the first both animals attacked the balls visually and took them readily (Figure 3). On day 2 they were also shown a clear rough ball and given shocks for taking it. At first they attacked it visually and took it under the web, but rapidly learned not to come out to it and by day 7 nearly always rejected it when it was placed against the arms” (Michels et al. 1987) Michels et al. 1987; Marini et al. 2017, Table 3 1 (On one specimen of bimaculatus, one of vulgaris) "Both animals were fed with crabs and pieces of shrimp on the subsequent days and by August 11th (=Day 1) were taking them readily and training was begun with smooth plastic black and while balls 12 mm in diameter. The octopuses were rewarded with a piece of shrimp for each attack at either ball, if it was taken under the web. The two colours were given alternately, each for either three or four trials a day. From the first both animals attacked the balls visually and took them readily (Figure 3). On day 2 they were also shown a clear rough ball and given shocks for taking it. At first they attacked it visually and took it under the web, but rapidly learned not to come out to it and by day 7 nearly always rejected it when it was placed against the arms” (Michels et al. 1987) Michels et al. 1987 1 (On one specimen of bimaculatus, one of vulgaris) "Both animals were fed with crabs and pieces of shrimp on the subsequent days and by August 11th (=Day 1) were taking them readily and training was begun with smooth plastic black and while balls 12 mm in diameter. The octopuses were rewarded with a piece of shrimp for each attack at either ball, if it was taken under the web. The two colours were given alternately, each for either three or four trials a day. From the first both animals attacked the balls visually and took them readily (Figure 3). On day 2 they were also shown a clear rough ball and given shocks for taking it. At first they attacked it visually and took it under the web, but rapidly learned not to come out to it and by day 7 nearly always rejected it when it was placed against the arms” (Michels et al. 1987) Michels et al. 1987 1 “The number of prey species included in an octopus’ diet was positively correlated with the time it resided in the same shelter. The inclusion of more species over time may simply result from chance encounters with rare species. However, it is also possible that when an octopus resides in one shelter for a long period of time, it learns the distributions of prey in the area and modifies its foraging accordingly…At Santa CataliNAIsland, most O. bimaculatus apparently stay in the same shelter, or at least the same area, for more than a month (Ambrose, 1982a). Since preferred prey species are generally rare, octopuses could enhance their exploitation of these species by learning their distributions in an area. This may explain the heavy utilization of chitons at Bird Rock. Most chitons were consumed by octopuses in one section of the study site. Since in general few chitons are found at the Bird Rock study side (Table III), it seems plausible that these octopuses had discovered an area of high chiton abundance, and were repeatedly exploiting it” (Ambrose 1984) Ambrose 1984 1 "Cephalopods also possess advanced spatial learning abilities, including exploratory learning and spatial maze learning (Mather, 1991; Forsythe & Hanlon, 1997; Boal et al., 2000a; Karson, Boal, & Hanlon, 2003; Jozet-Alves, Boal, & Dickel, 2008). For example, two-spot octopuses can use spatial cues to locate shelter in an unfamiliar arena. Most octopuses demonstrated spatial learning in a single day and were able to retain the information over seven days (Boal et al., 2000a)." (Schnell et al., 2020) Schnell et al., 2020 1 One specimen of O. bimaculatus: “Octopuses were conditioned to make visual attacks at a coloured smooth ball and also, by touch, to reject a transparent rough ball. They then showed no difference in rate of extinction of the visual responses to rough and smooth coloured balls. The long term memory ensuring such attacks was not impaired even by many negative contacts. The only weak sign of a cross modal influence was reduced tendency in some animals to make attacks four hours later. Thus the system has little or no capacity to learn by touch that a previously benign visual object has become harmful…In the tests with smooth balls given five hours later the proportion of attacks had recovered to about half of the original number at the beginning of the days and was similar for the balls that had been extinguished with rough and with smooth. Similarly the times taken to attack both coloured balls decreased in both animals to about half of those at the end of the extinction. There was no sign that the association of one colour with roughness and rejection had reduced its power to elicit a visual attack. By the following mornings the extinction had disappeared so completely that attacks at both balls were made at almost every trial. These two animals thus show no sign that the 77 contacts with the negative rough ball had influenced the positive visual memory” (Michels et al. 1987) Michels et al. 1987 One specimen of O. bimaculatus: “Octopuses were conditioned to make visual attacks at a coloured smooth ball and also, by touch, to reject a transparent rough ball. They then showed no difference in rate of extinction of the visual responses to rough and smooth coloured balls. The long term memory ensuring such attacks was not impaired even by many negative contacts. The only weak sign of a cross modal influence was reduced tendency in some animals to make attacks four hours later. Thus the system has little or no capacity to learn by touch that a previously benign visual object has become harmful…In the tests with smooth balls given five hours later the proportion of attacks had recovered to about half of the original number at the beginning of the days and was similar for the balls that had been extinguished with rough and with smooth. Similarly the times taken to attack both coloured balls decreased in both animals to about half of those at the end of the extinction. There was no sign that the association of one colour with roughness and rejection had reduced its power to elicit a visual attack. By the following mornings the extinction had disappeared so completely that attacks at both balls were made at almost every trial. These two animals thus show no sign that the 77 contacts with the negative rough ball had influenced the positive visual memory” (Michels et al. 1987) Michels et al. 1987 1 "Cephalopods also possess advanced spatial learning abilities, including exploratory learning and spatial maze learning (Mather, 1991; Forsythe & Hanlon, 1997; Boal et al., 2000a; Karson, Boal, & Hanlon, 2003; Jozet-Alves, Boal, & Dickel, 2008). For example, two-spot octopuses can use spatial cues to locate shelter in an unfamiliar arena. Most octopuses demonstrated spatial learning in a single day and were able to retain the information over seven days (Boal et al., 2000a)." (Schnell et al., 2020) Schnell et al., 2020 1 "Cephalopods also possess advanced spatial learning abilities, including exploratory learning and spatial maze learning (Mather, 1991; Forsythe & Hanlon, 1997; Boal et al., 2000a; Karson, Boal, & Hanlon, 2003; Jozet-Alves, Boal, & Dickel, 2008). For example, two-spot octopuses can use spatial cues to locate shelter in an unfamiliar arena. Most octopuses demonstrated spatial learning in a single day and were able to retain the information over seven days (Boal et al., 2000a)." (Schnell et al., 2020) Schnell et al. 2020 2 1 “The existence of a population comprised mainly of permanent residents has important implications for the ecology of Octopus bimaculatus. Mating and agonistic behavior may be influenced by familiarity between participants. A non-transient octopus population could result in a different pattern or intensity of resource utilization than a migratory or purely transient population.” (Ambrose 1982) & “The introduction of scorpionfish into a tank containing two or more octopuses appeared to stimulate fighting between the latter. This was perhaps competition for potential prey” (Taylor & Chen 1969) (Taylor & Chen 1969) 0 2007 in Bahía de Los Angeles, BC, Mexico: “females cease feeding after spawning because they cannot leave their eggs (Mangold 1987). In this study, 9 females were found caring for eggs; however, five had empty digestive tracts (and were eliminated from the analysis), and the remaining four ate mainly bivalves.” (Armendariz Villegas et al. 2014) (Armendariz Villegas et al. 2014) 1 2007 in Bahía de Los Angeles, BC, Mexico: “females cease feeding after spawning because they cannot leave their eggs (Mangold 1987). In this study, 9 females were found caring for eggs; however, five had empty digestive tracts (and were eliminated from the analysis), and the remaining four ate mainly bivalves.” (Armendariz Villegas et al. 2014) (Armendariz Villegas et al. 2014) 59 45 106.8 NY
Octopus salutii Octopus salutii 22 "This species occurs on tropical coral reefs from intertidal flats to at least 22 m deep" (Jereb et al., 2014) (Jereb et al., 2014; Mather et al., 2012) 0 "This species occurs on tropical coral reefs from intertidal flats to at least 22 m deep" (Jereb et al., 2014) (Jereb et al., 2014; Mather et al., 2012) 2 "The species inhabits the lower continental shelf and upper slope, primarily between 250 and 500 m depth" (Quetglas et al 2005) Quetglas et al 2005 15 50 "Mediterranean Sea and northeastern Atlantic. " (Jereb et al., 2014) (Jereb et al., 2014) 35 "Mediterranean Sea and northeastern Atlantic. " (Jereb et al., 2014) (Jereb et al., 2014) 1 730.5 Life span: 1.5-2 yrs (Mangold-Wirz et al., 1976) (Mangold-Wirz et al., 1976; Quetglas et al., 2005) 547.9 Life span: 1.5-2 yrs (Mangold-Wirz et al., 1976) (Mangold-Wirz et al., 1976; Quetglas et al., 2005) NA 0 NA 0 1 Present in the genus (Jereb et al., 2014) (Jereb et al., 2014) 33 "The stomach content analysis revealed a diet composed of 33 different prey items belonging to three major taxonomic groups (crustaceans, fishes and cephalopods). Quantitatively, crustaceans (Decapoda Reptantia and Natantia groups) were the most important prey, appearing in 87% of the stomachs, followed by fish (25%) and cephalopods (10%)." (Quetglas et al 2005) Quetglas et al 2005 0 1 Quetglas et al 2005 3 2 made up about 2 and 5% of cephalopod species found in stomachs of demersal sharks Scyliorhinus canicula and Squalus blainville caught in Aegean Sea (Kousteni et al. 2018) (Kousteni
et al. 2018)		 1 0 not gregarious (inferred from photo and video material) NA 0 NA 18 13 229.06 184.6 154.44 84 65 W-NY-W
Octopus vulgaris (all subspecies) Octopus vulgaris 800 "appeared in waters between 200 and 800 m depth" (Quetglas et al., 2000) (Quetglas et al., 2000; Sifner et al., 2005; Silva et al., 2011; Jereb et al., 2014; Lefkaditou et al. 1999; Quetglas et al 2005) 25 "Octopus salutii
caught throughout the year by bottom trawlers from 25
to 800 m depth in the western Mediterranean" (Quetglas et al 2005)		 (Quetglas et al., 2000; Sifner et al., 2005; Silva et al., 2011; Jereb et al., 2014; Lefkaditou et al. 1999; Quetglas et al 2005) 2 "benthic as adults" (Salman et al., 2003) Salman et al., 2003
Hanke & Kelber, 2020		 90 53 Records of up to 57 N in 1886, but rarely above 53 N (Rees, 1950) [Octopus vulgaris s.s.] (Rees, 1950) -37 25 S to 37 S (Jereb et al., 2014) [Octopus vulgaris type 3] (Jereb et al., 2014) 1 1 "It is known that O. vulgaris migrates to the coast during the first months of the year, and remains close to it (mainly at a depth between 30 and 60 m) during the reproductive period (Mangold-Wirz, 1963)." (Quetglas et al., 1998) (Elsevier, 2014; Quetglas et al., 1998; Alonso-Fernandez et al., 2017; Mereu et al., 2015; Villanueva & Norman, 2008; Pierce et al., 2010; Jereb et al., 2015; Boyle, 1983; García-Martínez et al., 2018; Alonso-Fernandez et al., 2017; Fadhlaoui-Zid et al., 2012; Brewer et al., 2017; Faraj & Bez, 2007; Jereb et al., 2014; Oosthuizen & Smale, 2003) 1 "Similarly,
Oosthuizen and Smale (2003) report that the population dynamics of O.
vulgaris in South Africa is likely dominated by the migration of ma-
turing females to deeper areas, in which they mature, spawn and brood," (Gonzalez-Gomez et al., 2020)		 (Pita et al., 2021; Zarrella et al., 2019; Ruby and Knudsen 1972; Mereu et al., 2015; Boyle, 1983; Alonso-Fernandez et al., 2017; Silva et al., 2002; Jereb et al., 2014; Gonzalez-Gomez et al., 2020) 0 "Some authors suggest that O. vulgaris seems to undergo seasonal
movements mainly vertically oriented, at different times, in relationship
to the size and/or maturity condition, and that there are animals that
probably do not migrate, like those which spawn in deeper waters or
stay near the coast during the winter (e.g., Guerra, 1975; Hatanaka,
1979; Mangold, 1983; Mangold-Wirz, 1963). However, when not
travelling in/offshore, O. vulgaris seems to be a truly sedentary species
(Mangold, 1983). There are only a few records of horizontal movements. Itami (1964) in the Japan Sea branded 1559 O. vulgaris, 14 of which were
recovered within two months and the longest straight distance from
release to recapture was 48 km (see Nagasawa et al., 1993). In the
Mediterranean Sea the preliminary data on movements were obtained
from 9 recaptured specimens in the same area investigated in the
present study (central-western Sardinian Sea). The Authors reported
1.2 km as the longest straight distance travelled by five males (2 mature
and 3 spawning specimens), after 8 and 30 days of freedom, respectively
(Mereu et al., 2010)." (Mereu et al., 2015)		 (Mereu et al., 2015) 1461 "Mangold-Wirz (1963) gave careful consideration to the age and lifespan of the
Mediterranean cephalopods. Her conclusions were based upon the excellent cross-
sectional data of growth that she had collected. Sexual maturation in O. vulgaris
occurred when the dorsal mantle length of the males was 80 mm at about 8-10
months of age, and in females with a dorsal mantle length of 130-140 mm at
18-24 months of age (Table 1). Shorter periods were also tentatively suggested, 7
months for the males and 15-17 months for the females. The maximum age was
given as 2 years, possibly 3-4 years, for the male, and 2 or possibly 3 years for the
female." (Nixon, 1969)		 (Nixon, 1969; Casalini et al., 2020; Quinteiro et al., 2020; De Luca et al., 2016; Pita et al., 2021; Villanueva & Norman, 2008; Silva et al., 2019; Miliou et al 2006; Mauvisseau et al., 2017; Lima et al., 2017; Garci et al., 2016; Lourenco et al., 2015; Moreno et al., 2014; Kivengea et al., 2014; Fuentes & Iglesias, 2010; Jereb et al., 2015; Pierce et al., 2010; Boyle, 1983; Nixon, 1969; Jereb et al., 2014; Perales-Raya et al., 2014; Perales-Raya et al., 2014; Cuccu et al., 2013; Lourenco et al., 2012; Fadhlaoui-Zid et al., 2012; Fuentes & Iglesias, 2010; Fuentes & Iglesias, 2010; Katsanevakis & Verriopoulos, 2006; Jereb et al., 2015; Hernández-López & Castro-Hernández, 2001; Domain et al., 2000; Raya & Hernandez-Gonzalez, 1998; Jereb et al., 2015; Walter González et al., 2015; Castanhari & Gomas Tomas, 2012; Jereb et al., 2014; Fadhlaoui-Zid et al., 2012; Jereb et al., 2015; Hernández-López & Castro-Hernández, 2001; Smale & Buchan, 1981; Hernández-López & Castro-Hernández, 2001; Hanke & Kelber, 2020; Nafkha et al., 2019; Jimenez-Prada et al., 2014; Perales-Raya et al., 2014; Hernández-López & Castro-Hernández, 2001; Canali et al., 2011) 60.9 [Notice the small individuals in the sample] "Although this study
was restricted to a particular range of BW (average
350 g), the observed number of rings showed a quite
remarkable range (between 72 and 371 increments).
On the basis of assumed daily deposition, this sug-
gests an estimated age of ~2 to 12 mo for the sampled
specimens. " (Canali et al., 2011)		 (Nixon, 1969; Casalini et al., 2020; Quinteiro et al., 2020; De Luca et al., 2016; Pita et al., 2021; Villanueva & Norman, 2008; Silva et al., 2019; Miliou et al 2006; Mauvisseau et al., 2017; Lima et al., 2017; Garci et al., 2016; Lourenco et al., 2015; Moreno et al., 2014; Kivengea et al., 2014; Fuentes & Iglesias, 2010; Jereb et al., 2015; Pierce et al., 2010; Boyle, 1983; Nixon, 1969; Jereb et al., 2014; Perales-Raya et al., 2014; Perales-Raya et al., 2014; Cuccu et al., 2013; Lourenco et al., 2012; Fadhlaoui-Zid et al., 2012; Fuentes & Iglesias, 2010; Fuentes & Iglesias, 2010; Katsanevakis & Verriopoulos, 2006; Jereb et al., 2015; Hernández-López & Castro-Hernández, 2001; Domain et al., 2000; Raya & Hernandez-Gonzalez, 1998; Jereb et al., 2015; Walter González et al., 2015; Castanhari & Gomas Tomas, 2012; Jereb et al., 2014; Fadhlaoui-Zid et al., 2012; Jereb et al., 2015; Hernández-López & Castro-Hernández, 2001; Smale & Buchan, 1981; Hernández-López & Castro-Hernández, 2001; Hanke & Kelber, 2020; Nafkha et al., 2019; Jimenez-Prada et al., 2014; Perales-Raya et al., 2014; Hernández-López & Castro-Hernández, 2001; Canali et al., 2011) 1461 "Mangold-Wirz (1963) gave careful consideration to the age and lifespan of the
Mediterranean cephalopods. Her conclusions were based upon the excellent cross-
sectional data of growth that she had collected. Sexual maturation in O. vulgaris
occurred when the dorsal mantle length of the males was 80 mm at about 8-10
months of age, and in females with a dorsal mantle length of 130-140 mm at
18-24 months of age (Table 1). Shorter periods were also tentatively suggested, 7
months for the males and 15-17 months for the females. The maximum age was
given as 2 years, possibly 3-4 years, for the male, and 2 or possibly 3 years for the
female."" (Nixon, 1969)		 (Boyle, 1983; Nixon, 1969; Cuccu et al., 2013; Perales-Raya et al., 2014; Jereb et al., 2015; Jimenez-Prada et al., 2014; Smale & Buchan, 1981) 60.9 "Males become mature at a smaller size and a younger age than females; the testis increases in relative size probably 2-3 months before the ovary begins to enlarge. In the Catalonian Sea. males weighing 200g are mostly mature. while the smallest females with ripe eggs have a body weight of about 500 g. In females. the ovary. the oviducal glands and the oviducts enlarge enormously about 3 to 4 months before spawning" (Boyle, 1983). (Boyle, 1983; Nixon, 1969; Cuccu et al., 2013; Perales-Raya et al., 2014; Jereb et al., 2015; Jimenez-Prada et al., 2014; Smale & Buchan, 1981) 8 8 1 "Octopus vulgaris [is] an ambush predator, hiding behind stones to swoop out and attack a passing crab" (Hanlon & Messenger, 2018, p. 82). Hanlon & Messenger, 2018, p. 82
Bennice et al., 2021		 1 "As a bottom feeder, foraging often seems to be tactile (Jereb et al., 2014), involving exploration of the surroundings with its arms, in search for crustaceans, fish, shelled molluscs or polychaetes (Mather, 1991; Boyle and Rodhouse, 2005; Mather et al., 2012; Sanchez et al., 2015). In addition, visual and chemical cues are most likely used to find prey ( Boyle and Rodhouse, 2005)." (Hanke & Kelber, 2020) Hanke & Kelber, 2020
Jereb et al., 2014
Bennice et al., 2021
Hernandez-Urcera et al., 2019
Gilchrist 2003
Mather, 1991		 1 "Another dynamic pattern used for hunting is the chromatic pulse of Octopus vulgaris (Packard and Hochberg 1977; Mather and Mather 2004). This pattern may deceive the prey by simulating an approaching object and inducing the prey to move closer to the predator (How et al. 2017)." (Carreno-Castilla et al 2020) Carreno-Castilla et al 2020
Packard and Sanders (1969)
Hanlon & Messenger, 2018, p. 80-81		 1 "Despite their flexibility, octopuses preferentially use chemotactile methods to hunt for prey in rock and coral crevices while hovering through the seabed (speculative hunting), while the visual senses are primed for predatory avoidance (camouflage matching) and ambushing prey that wanders close to their den (Mather & O’Dor 1991, Forsythe & Hanlon 1997)." (Sampaio et al., 2018) Sampaio et al., 2018
Cosgrove, 2002
Hanlon & Messenger, 2018, p. 81
Iglesias et al., 2014
FIorito & Gherardi, 1999
Jereb et al., 2014
Bennice et al., 2021
Gilchrist 2003
Mather & O´Dor, 1991
Mather, 1991		 1 "Whilst SCUBA diving during a routine fish transect in the mooring gulley at Storm's River Mouth (grid 3423BB) in the Tsi- tsikama Coastal National Park, one of us (CB) observed a medium-sized Hawksbill Turtle lEretmochelys imbricata)" apparently lodged under a rock overhang. Closer observation revealed that the turtle was restrained by a large octopus that was feeding on the left hind flipper. The incident occurred at 4 p.m. on 28 April 1983. Underwater visibility was good and the water temperature 18°C. The mooring gulley is formed by east-west running sandstone ridges that break the force of wave action. Water depth was 4m." (Buxton & Branch, 1983) Buxton & Branch, 1983 1 "The two most frequently used behaviors during foraging events for both species were speculative bottom searching and sitting. The next three most frequently used behaviors (in decreasing order) for O. vulgaris were grope searching, parachute attack, and crawling and for M. defilippi were flounder mimicry swimming, tripod stance, and grope searching. " (Bennice et al., 2021) [Location South Florida lagoon] Gilchrist 2003 1 "It is well known that O. vulgaris paralyzes different species of crabs injecting cephalotoxin prior to ingestion (Ghiretti, 1960)" (Hernández-Urcera et al., 2014) Hernández-Urcera et al., 2014
Iglesias et al 2014
Hiemstra, 2015
Arnold &
Arnold, 1969; Nixon, 1980
Anderson et al., 2008
Roura et al., 2010
Jereb et al., 2015
Hernandez-Garcia et al., 2000
FIorito & Gherardi, 1999
Hanlon & Messenger, 2018, p. 93
Nixon & Maconnachie, 1988
Wodinsky, 1978
FIorito & Gherardi, 1999
Guerra et al., 2014
McQuaid, 1994
Nixon & Maconnachie, 1988
Niven, 1988
Guerra & Nixon, 1987
Jereb et al., 2014
Blustein & Anderson, 2016
Gilchrist 2003
Niven, 1988		 1 " Regardless of their prey size, Octopus will always try the pulling method first (Fiorito & Gherardi, 1999)" [in regards to mullos] (Hiemstra, 2015) Hiemstra, 2015
Hanlon & Messenger, 2018, p. 94
FIorito & Gherardi, 1999
Guerra et al., 2014
Mather, 1991
McQuaid, 1994
Jereb et al., 2014		 Guerra et al., 2014 11 10 1 "Den use (Mereu et al., 2014)" Mereu et al., 2014
Guerra et al., 2014)
Hanke & Kelber, 2020
Katsanevakis and Verriopoulos2004
Edmunds1974		 1 Octopus inhabiting a shallow, small-boulder substratum made extensive modifications to their habitat, excavating dens of up to 1 m deep in the sand below the boulders. These dens were not visible during the day as the octopus appeared to retract the small boulders over their den entrances. This unique behavioural strategy is thought to be a means to reduce predation and reduce light intensity during the day. Octopus were not observed in the small-boulder habitat during the five hours of daytime sampling. With nocturnal activity and extensive habitat modification, it is likely that avoidance of predation may be an important driver influencing the behaviour of the octopus population under study. " (De Beer & Potts, 2013) De Beer & Potts, 2013
Villanueva & Norman, 2008
Niven, 1988
Mathers 1984
Alves et al., 2008
Niven, 1988		 1 "The common octopus is said to be nocturnal (Woods, 1965; Altman, 1966; Kayes, 1974; Jereb et al., 2014), but it has been seen to shift its activity phase, for example in the presence of prey or predators (Meisel et al., 2013), and thus some studies report crepuscular or even diurnal activity (Mather, 1988; Meisel et al., 2003, 2006)." (Hanke & Kelber, 2020) Hanke & Kelber, 2020
Mather, 2019		 1 "This seems particularly likely given the octopuses often consume food at a den or other shelter (Mather 1991a), and spend the majority of their time hiding (e.g. O. vulgaris: Mather 1988, E. dofleini: Scheel and Bisson 2012), which allows time to drill, chip, or pull open prey in safety (see Sect. 2.2)" (Iglesias et al., 2014). Iglesias et al., 2014
Niven, 1988
Mather, 1991		 1 "Despite their flexibility, octopuses preferentially use chemotactile methods to hunt for prey in rock and coral crevices while hovering through the seabed (speculative hunting), while the visual senses are primed for predatory avoidance (camouflage matching) and ambushing prey that wanders close to their den (Mather & O’Dor 1991, Forsythe & Hanlon 1997)." (Sampaio et al., 2018) Sampaio et al., 2018
Mather & Mather, 1994
Josef et al., 2012
Hanlon & Messenger, 2018, p. 111		 1 During the prey capture sequence, a change in chromatophore pattern usually takes place between the second and third phase (hernández-García et al. 2000). During the attention phase, the para larvae maintain contracted chromatophores, so that the octopus is nearly transparent to a human observer; then, during the positioning phase and/or during the contact with the prey, the chromatophores from the dorsal mantle, head and arms are expanded dramatically. After seizure of the prey the chromatophores are contracted again. This visual signal is suspected to be defensive behaviour (see ‘Defences’, p. 168). selection of the attack angle probably depends on the type of prey, its size and defences (i.e., many crustacean zoeae possess dorsal spines) (Figure 28). " (Villanueva & Norman, 2008) Villanueva & Norman, 2008
Hernandez-Garcia et al., 2000
Hanlon & Messenger, 2018, p. 116		 1 During the prey capture sequence, a change in chromatophore pattern usually takes place between the second and third phase (hernández-García et al. 2000). During the attention phase, the para larvae maintain contracted chromatophores, so that the octopus is nearly transparent to a human observer; then, during the positioning phase and/or during the contact with the prey, the chromatophores from the dorsal mantle, head and arms are expanded dramatically. After seizure of the prey the chromatophores are contracted again. This visual signal is suspected to be defensive behaviour (see ‘Defences’, p. 168). selection of the attack angle probably depends on the type of prey, its size and defences (i.e., many crustacean zoeae possess dorsal spines) (Figure 28). " (Villanueva & Norman, 2008) Villanueva & Norman, 2008 1 Defensive tool use: "Other remarkable anti-predatory strategies reported among octopuses include defensive tool-use (Fig. 2). A common octopus was recently featured in the BBC Blue Planet II series using its suckered arms to gather and create a protective armour of shells and stones to protect itself from hunting sharks (Jeffs & Brownlow, 2017)." (Schnell et al., 2020) Schnell et al., 2020
Ambrose, 1983
Mather, 1994
Niven, 1988		 1 "In the Atlantic waters of Spain, Hernández- Urcera et al. (2020) observed a small- sized O. vulgaris (sensu stricto) performing a de- fense behavior that has been classified as bipedal locomotion" (Amodio et al., 2021) Amodio et al., 2021
Soto et al., 2020
Hanlon & Messenger, 2018, p. 125		 1 Inking multiple times (Hernandez-Urcera et al., 2019) Hernandez-Urcera et al., 2019
Niven, 1988		 1 (Jereb et al., 2014) 1 Here we see rapid neural polyphenism being exercised at its best. The cognitive aspects of these complex behaviours deserve future study because, while rapidly fleeing, the octopuses were tailoring their next behaviours to the benthic surroundings. An example is the ‘fake right, go left’ behaviour seen often in these trials (Fig. 5.30a) in which the swiftly swimming octopus suddenly stops, inks to the right, blanches and descends left amidst a patch of mottled algae (all in 0.4 seconds), and matches the mottled algal background. Obviously there is still much yet to learn about secondary defences in octopuses and how their decision-making is so precise." (Hanlon & Messenger, 2018, p. 141-143) Hanlon & Messenger, 2018, p. 141-143
Boyle, 1983		 193 "O. vulgaris fed basically on crustaceans (mainly decapods) and fishes, although it occasionally included gastropods and cephalopods in its diet." (Quetglas et al., 1998) [Location Mediterranean] ["The importance of fishes was higher when compared to the values obtained by Guerra (1978), Smale and Buchan (1981) and SaÂnchez and Obarti (1993), but similar to those found by Nigmatullin and Ostapenko (1976)."] Sakamoto et al., 2006
Niven, 1988
Nigmatullin and Ostapenko (1976)
Quetglas et al 1998
Pierce et al 2010
Nixon & Budelmann, 1984
Ulas et al., 2019
Abel & Nivel, 1990
Diaz et al., 2005
Tirelli et al., 2000
Nixon & Maconnachie, 1988
Kuhlmann & McCabe, 2014
Jereb et al., 2014		 1 (Hanlon & Messenger, 2018, p. 87) 1 Quetlag et al. (1998)
Jereb et al., 2014		 8 32 "Coastal fish (Epinephelus marginatus, Serranus sp., AtheriNApresbyter) attracted to O. vul-
garis egg masses during hatching periods have been observed preying on paralarvae
(Villanueva and Norman, 2008). Further, paralarvae of 6.5–18 mm TL have been rec-
orded in the stomach contents of albacore (Thunnus alalunga) (Bouxin and Legendre,
1936). Predators of subadult and adult O. vulgaris include fish, marine mammals, birds, man,
and other cephalopod species (Hanlon and Messenger, 1996). Octopus vulgaris has been
found in the stomachs of bottlenose dolphin (Tursiops truncatus) (Blanco et al., 2001),
Risso’s dolphin (Grampus griseus) (Blanco et al., 2006), and Mediterranean monk seal
(Monachus monachus) (Pierce et al., 2011) in the Mediterranean Sea. Marine mammal
predators of O. vulgaris in Galician waters include common dolphin (Delphinus delphis),
long-finned pilot whale (Globicephala melas), and sperm whale (Physeter macrocephalus)
(Table 3.3, see also at González et al., 1994a; López, 2002; Santos et al., 2004a, 2013, 2014). " (Jereb et al., 2015)		 (Mather, 1991) (Meisel et al., 2013) (Pierce et al., 2010) (Boyle,
1983) (Niven, 1988) (Jereb et al., 2014) (Karamanlidis et al., 2020) (Di
Lorenzo et al., 2020) (Tonay et al., 2016) (Quigley & Flannery 2014) (Villanueva
& Norman, 2008) (Matic-Skoko et al., 2014) (Garibaldi & Orsi Relini,
2012) (Abel & Nivel, 1990) (Jereb et al., 2015) (Gilchrist, 2003) (George-Nascimento
et al., 1985) (De Beer & Potts, 2013)		 5 1 "Octopuses typically avoid each other. In O. vulgaris, Altman (1967) observed dens close together but there were no signs of interactions between neighbors." (Alves et al., 2008). "Octopus vulgaris is solitary, and the sexes only meet during mating (Hanlon and Messenger, 2018)" (Hanke & Kelber, 2020) Alves et al., 2008; Hanke & Kelber 2020 0 "Octopuses typically avoid each other. In O. vulgaris, Altman (1967) observed dens close together but there were no signs of interactions between neighbors." (Alves et al., 2008). "Octopus vulgaris is solitary, and the sexes only meet during mating (Hanlon and Messenger, 2018)" (Hanke & Kelber, 2020) Alves et al., 2008; Hanke & Kelber 2020 23 18 1 "Kuba, Byrne, Meisel & Mather (2006a) have studied it, together with habituation, in O. vulgaris. When an item (a miniature plastic crab) is moved outside the octopus tank, it quickly habituates both within and across trials." Mather 2019 1 Table 3 in 'Behavioral analysis of learning and memory in Cephalopods' Borelli & Fiorito in Byrne eds. 2008 Borelli & Fiorito in Byrne eds. 2008 1 Table 3 in 'Behavioral analysis of learning and memory in Cephalopods' Borelli & Fiorito in Byrne eds. 2008 Borelli & Fiorito in Byrne eds. 2008 1 "Successful performance in the task requires the learning of the association between interacting with the maze and receiving a food reward. The octopus never sees the location of the reward, nor does it see its arm. Yet it learns to insert an arm through the maze to a goal box to retrieve it, thus associating the action of the arm with the subsequent reward. It has been shown that in cephalopods, positive reinforcement learning using food as a reward is processed in the brain [1, 7]. This leads us to conclude that learning of the tactile and proprioceptive tasks was mediated by the CNS and could not be solely achieved by the arms alone." (Gutnick et al., 2020) Gutnick et al., 2020 1 "Octopuses learn to not respond to (i.e. not attack) a stimulus that is always associated with a negative reinforcement (electric shock), known as passive avoidance learning" (Borelli et al. 2020) Borrelli et al. 2020 1 "Conditional discrimination in the octopus (Octopus vulgaris) was studied using successive discrimination training. The experimental animals were divided into two groups, and a barrel-shaped white object (stimulus) was presented to each group. One of the groups was rewarded with food for responding to the stimulus, but only when the tank was aerated, whereas the other group was rewarded with food for responding to the stimulus when the aeration was switched off. The number of trials in which octopuses responded to the stimulus, and the latency of the responses, were significantly different between trials with the aeration on and trials with the aeration off, in both groups. Therefore, the octopuses learned to conditionally discriminate." Tokuda et al. 2015 "Only one octopus learnt the replenishing rates of different prey types and was able to use these rules to solve an episodic-like memory task. When analysing the strategies used by tested octopuses during the replenishing rate training, such as familiarity, risk proneness, spontaneous alternation and win–stay, we observed above all a high interindividual variability.Only one octopus learnt the replenishing rates of different prey types and was able to use these rules to solve an episodic-like memory task. When analysing the strategies used by tested octopuses during the replenishing rate training, such as familiarity, risk proneness, spontaneous alternation and win–stay, we observed above all a high interindividual variability….One individual (Teddy) learnt the replenishing rate of the different food items and subsequently succeeded in the episodic-like memory task. This may indicate that O. vulgaris possesses the neural prerequisites for episodic-like memory." (Poncet et al. 2022) Poncet et al. 2022 1 "In Mackintosh & Mackintosh ( I 963), eight octopuses were trained to discriminate
between black and white vertical rectangles in order to study reversal learning and the
role of irrelevant cues. Only the initial task will be discussed here. Partitions similar to
those used by Sutherland et al. (1963), were used to separate the two stimuli and to
ensure a central start position. No mention was made of a transverse bar being rotated
to introduce stimuli ; I assume objects were supported by hand. Subjects were trained
until they reached a criterion of 90% success over 2 d of 10 trials each. Only average
errors to criterion were published. These were so few (2.67) that animals must have
reached criterion almost immediately. " (Boal, 1996)		 Boal, 1996 1 Table 3 in 'Behavioral analysis of learning and memory in Cephalopods' Borelli & Fiorito in Byrne eds. 2008 Borelli & Fiorito in Byrne eds. 2008 1 "O. vulgaris learned to direct an arm in a maze on the basis of visual cues (Gutnick, Byrne, Hochner & Kuba, 2011), or kinesthetic or tactile ones (Kuba, personal communication)" (Mather, 2019) Mather 2019 1 "O. vulgaris learned to direct an arm in a maze on the basis of visual cues (Gutnick, Byrne, Hochner & Kuba, 2011), or kinesthetic or tactile ones (Kuba, personal communication)" (Mather, 2019) Mather 2019 0 "Octopuses did not succeed in recognizing themselves in even a partial test
of Gallup’s (1995) mirror self-recognition task (Mather, Carere, Fiorito & Anderson, 2018). O. vulgaris perceived this visual feedback as an anomalous situation and not a view of a conspecific; they made more mantle-up challenge displays to conspecifics and more Passing Cloud displays to the mirror." (Mather, 2019)		 Mather 2019 1 "A recent study shows that avoidance conditioning in the cephalopod Octopus vulgaris is mediated by long-term potentiation (LTP), a form of synaptic plasticity thought to be important in vertebrate associative learning. Thus, LTP appears to be an evolutionarily conserved learning mechanism." "These results, as well as implicating LTP in avoidance learning in the octopus, demonstrate a clear segregation of short-term and long-term memory systems in the brain of this animal. Short-term memories are stored within the neural circuits that produce the attack behavior, whereas consolidated memories are stored in the vertical lobe (at least for the first 24 hours). In its separation of the sites of short-term and long-term memory storage, the brain of the octopus resembles those of vertebrates [11]." (Glanzmen, 2008) Glanzmen, 2008 1 "A recent study shows that avoidance conditioning in the cephalopod Octopus vulgaris is mediated by long-term potentiation (LTP), a form of synaptic plasticity thought to be important in vertebrate associative learning. Thus, LTP appears to be an evolutionarily conserved learning mechanism." "These results, as well as implicating LTP in avoidance learning in the octopus, demonstrate a clear segregation of short-term and long-term memory systems in the brain of this animal. Short-term memories are stored within the neural circuits that produce the attack behavior, whereas consolidated memories are stored in the vertical lobe (at least for the first 24 hours). In its separation of the sites of short-term and long-term memory storage, the brain of the octopus resembles those of vertebrates [11]." (Glanzmen, 2008) Glanzmen, 2008 1 "Further investigation of play-like behavior suggested it is a wider phenomenon in octopuses. Kuba et al. (2006) tested Octopus vulgaris Cuvier, 1797 with presentations of plastic blocks, clam prey, and empty clam shells. The octo- puses ate the clams, ignored the empty shells, and sometimes engaged in play-like behavior with the blocks. The play be- havior consisted of passing the block from arm to arm, extending the arm and pulling it back near the body, and pulling the block along as the octopus moved. These were designated only play-like on the basis of numbers of repeti- tions. The arms were clearly central to the actions, and the peak of playful behavior came during the sixth of ten trials, after which the octopuses habituated again. Interestingly, Kuba et al. (2006) found that, unlike in mammals (Fagen 1981, Power 2000), young and adult octopuses played the same amount. For the solitary octopus, play was not the result of needing to learn the nuances of a social group, nor was it restricted to the protected environment of the family. It did not occur frequently but occurred equally at different times in the lifespan." (Mather, 2008) Mather 2008 1 Tool use: "the octopuses also used water jetting in manipulation tasks, such as cleaning out items or sites
used as ‘homes’ and repelling scavenging fish (Mather, 1992)" (Mather, 2019)		 Mather 2019 1 "Play is much more common in young mammals than adults and is ‘functional’ as practice for future adaptive behaviours (Burghardt, 2005). Like the keas (Diamond & Bond, 2004), subadult octopuses in the Kuba et al. (2006b) study did not show more play-like behavior than mature adults. Play might occur when complex animals with a heavy dependence on learning have excess resources and a limited environment and are ‘bored’ (see discussion by Kuba et al., 2014). Inglis et al. (2001) has pointed out, however, that foraging animals in a complex and varying environment make a tradeoff between immediate use of resources and information acquisition for future use. They explore more when they are satiated and safe, as in the Mather & Anderson (1993) play situation. This tradeoff between present and potential future rewards is also described as ‘latent learning’ and ‘contra-freeloading’." (Mather, 2019) Mather 2019 1 "Laboratory studies by Schiller (1949) and Wells
(1964) tested the abilities of O. vulgaris to solve a detour
task and explored the strategies the octopuses used. In both
experiments, octopuses were trained to detour around
opaque partitions to reach a crab visible behind a trans-
parent wall, but not directly accessible to them (Fig. 1). In
Wells’ experiment (1964), only eight octopuses out of 29
succeeded in solving the detour problem in the first trial.
The failure of the majority of the octopuses could be
interpreted as demonstrating an inability to show detour
behavior, and hence an absence of spatial representation in
octopuses. However, Regolin et al. (1994) showed that in chicks, performances in detour problems can be affected by
perceptual and motivational factors. In the experiments of
Schiller (1949) and Wells (1964), the transparent wall that
separated the octopuses from the crabs could have been
such an unnatural stimulus that the animals failed to per-
ceive it as an obstacle. With repeated trials, the octopuses
showed some improvement in performance, spending less
time attacking through the glass before entering the central
alley, and all animals learned to complete the task. But
what did the octopuses learn? Schiller (1949) suggested
that octopuses needed to maintain constant tactile contact
with the wall separating them from the goal. Wells later
showed (1964) that if the tactile contact was lost, the
octopuses needed to maintain a continuous visual fixation
on the wall. Thus, detour behavior in octopuses was visu-
ally guided; bodily position was not used to compute
position of the goal relative to their body. These detour
experiments did not provide clear evidence for spatial
representation in octopuses; however, they did show that
octopuses can develop an efficient strategy to solve a
spatial task." (Alves et al., 2008)		 Alves et al. 2008 1 "O. vulgaris learned to direct an arm in a maze on the basis of visual cues (Gutnick, Byrne, Hochner & Kuba, 2011), or kinesthetic or tactile ones (Kuba,
personal communication)" (Mather, 2019)		 Mather 2019 1 “O. vulgaris may split the appearance of its body in two halves (generally left and right) in which the animal <<may…be in a different phase on one side of the body from that on the other: one side dark, the other light>> [Packard & Sanders (1969, p. 93)]” (Borelli et al. 2006) Borelli et al. 2006 4 1 "This study reports the first experimental evidence of O. vulgaris’ ability to recognise a familiar conspecific and to remember it for at least one day. As shown during the test phase, unfamiliar pairs, i.e. pairs composed of individuals that have had no previous experience of each other, executed more numerous physical contacts and showed shorter latencies than familiar pairs, being thus more aggressive and prone to interact. Besides, reversals of dominance (i.e. alphas switched to betas and, consequently, betas to alphas) were only observed in unfamiliar pairs. Taken together, these results seem to support our hypothesis that O. vulgaris can discriminate familiar from unfamiliar conspecifics, meaning that it is able of, at least, class-level or binary IR sensu [10]." (Tricarico et al., 2011) Tricarico et al., 2011 0 "Octopuses typically avoid each other. In O.
vulgaris, Altman (1967) observed dens close together but
there were no signs of interactions between neighbors." (Alves et al., 2008)		 (Alves et al., 2008) 1 In lab "This study reports the first experimental evidence of O. vulgaris’
ability to recognise a familiar conspecific and to remember it for at
least one day. As shown during the test phase, unfamiliar pairs, i.e.
pairs composed of individuals that have had no previous
experience of each other, executed more numerous physical
contacts and showed shorter latencies than familiar pairs, being
thus more aggressive and prone to interact. Besides, reversals of
dominance (i.e. alphas switched to betas and, consequently, betas
to alphas) were only observed in unfamiliar pairs. Taken together,
these results seem to support our hypothesis that O. vulgaris can
discriminate familiar from unfamiliar conspecifics, meaning that it is able of, at least, class-level or binary IR sensu [10]." (Tricarico et al., 2011)		 (Tricarico et al., 2011) 1 "Spawning took place over the whole continental shelf up to 100 m deep, with sometimes more spawning close to the coast, whereas recruitment seemed systematically to be concentrated at the coast, shallower than 50 m. Further, the area used for spawning was highly
variable annually throughout the study period between 1998 and
2003, whereas the areas of recruitment were more stable." (Faraj & Bez, 2007) but see "aggregations have been observed in O. vulgaris pre-recruits in the Portuguese coast (Moreno et al., 2014)." (Puerta et al., 2016)		 (Faraj & Bez, 2007) 1 “(1) As soon as the female is put in at the far end of the tank, about 50 cm away, the male emerges from his home of bricks. His colour is variable; he may flush darkly, very often flattening and spreading the web, thus appearing much larger than normal; or, more rarely, he pales and spreads out, producing the almost white 'dymantic' colouration (Boycott & Young 1950). At this stage the female is typically quiet, sitting on the side or bottom of the tank. (2) The male approaches the female with one or more arms extended or makes a jet-propelled rush and pins her down by covering her with his web, the latter becoming more likely when the male has been in the tank for a matter of weeks rather than days. The reddish/brown colour of the male is darkest at this point; the female is typically much paler. (3) Immediately this contact with the female has been made, the male begins to probe with the third right arm (see Table I) . (4) Usually the female struggles very little once the male (in nearly all our experiments somewhat the larger of the two) has pinned her down in his first enveloping rush. In this event the male may get his hectocotylus into the mantle cavity and begin to arch and pump within 1 min of his first sight of the female. (5) Successive spermatophores are passed at intervals of about 15 s, and copulation may last for 1 hr or more (Racovitza 1894). We normally separated the animals after two or three arching and pumping” (Wells & Wells 1972) (Wells & Wells 1972) 0 " The female stays with the eggs for the duration of development, which can last up to 5 months, continuously caring for and defending the eggs. The female octopus does not feed during this period, digesting its own musculature in this last phase of its life (Jereb et  al., 2014;
Hanlon and Messenger, 2018)." (Hanke & Kelber, 2020)		 (Hanke & Kelber, 2020) & (Anderson et al., 2002) & (Jereb et al., 2014) 1 " The female stays with the eggs for the duration of development, which can last up to 5 months, continuously caring for and defending the eggs. The female octopus does not feed during this period, digesting its own musculature in this last phase of its life (Jereb et  al., 2014;
Hanlon and Messenger, 2018)." (Hanke & Kelber, 2020)		 (Hanke & Kelber, 2020) & (Anderson et al., 2002) & (Jereb et al., 2014) 1561 1253 538.572 239.316 171.6 155 80 80 W-W-NY
Ocythoe tuberculata Ocythoe tuberculata 317 [AL: Has been caught at greater depths but no evidence for adult specimens] "B ATHYMETRIC DISTRIBUTION .—The species occupied a wide bathymetric range from 17
to 317 m, including coastal waters without river outflow and the intertidal zone. The most
coastal specimen fished measured 20 cm ML and weighed 2900 g BW. The deepest speci-
men caught measured 18 cm ML and weighed 1864 g BW." (Silva et al., 2002)		 (Salman et al., 2003; Cairns, 1976; Robson, 1929; Haimovici et al., 1989; Quetglas et al., 2000; Sifner et al., 2005; Silva et al., 2011; Zarrella et al., 2019; Ruby and Knudsen 1972; Mauvisseau et al., 2017; Lima et al., 2017; Morene et al., 2014; Pierce et al., 2010; Katsanevakis & Verriopoulos, 2004; Boyle, 1983; Elsevier, 2014; Boyle & Rodhouse, 2005; Jereb et al., 2014; Pita et al., 2021; Sanchez & Martin 1993; Roditi et al., 2020; Ulas et al., 2019; Perales-Raya et al., 2018; Banon et al., 2018; Garci et al., 2016; Guerra et al., 2015; Morene et al., 2014; Mayo-Hernandez et al., 2013; Garofalo et al., 2010; Carreira & Goncalves, 2009; Silva et al., 2002; Jereb et al., 2014; Hernandez-Urcera et al., 2019; Sillero-Rios et al., 2018; Flores-Valle et al., 2018; Avendano et al., 2020; Haimovici & Andriguetto, 1986; Wittmann & Griffiths, 2017; Mereu et al., 2015; Gonzalez-Gomez et al., 2020; Ulas et al., 2011; Mather et al., 2012) 0 0-200 m (Haimovici et al., 1989) (Salman et al., 2003; Cairns, 1976; Robson, 1929; Haimovici et al., 1989; Quetglas et al., 2000; Sifner et al., 2005; Silva et al., 2011; Zarrella et al., 2019; Ruby and Knudsen 1972; Mauvisseau et al., 2017; Lima et al., 2017; Morene et al., 2014; Pierce et al., 2010; Katsanevakis & Verriopoulos, 2004; Boyle, 1983; Elsevier, 2014; Boyle & Rodhouse, 2005; Jereb et al., 2014; Pita et al., 2021; Sanchez & Martin 1993; Roditi et al., 2020; Ulas et al., 2019; Perales-Raya et al., 2018; Banon et al., 2018; Garci et al., 2016; Guerra et al., 2015; Morene et al., 2014; Mayo-Hernandez et al., 2013; Garofalo et al., 2010; Carreira & Goncalves, 2009; Silva et al., 2002; Jereb et al., 2014; Hernandez-Urcera et al., 2019; Sillero-Rios et al., 2018; Flores-Valle et al., 2018; Avendano et al., 2020; Haimovici & Andriguetto, 1986; Wittmann & Griffiths, 2017; Mereu et al., 2015; Gonzalez-Gomez et al., 2020; Ulas et al., 2011; Mather et al., 2012) 1 pelagic octopod (Tejerina, 2019) Tejerina, 2019 122 62 62.5 N to 60 S (Jereb et al., 2014) (Jereb et al., 2014) -60 62.5 N to 60 S (Jereb et al., 2014) (Jereb et al., 2014) 1 1 "The occurrence of O. tuberculata is often explained related to sea currents and sea warming. In the Mediterranean Sea (type locality), this species is probably transported from the open waters of the Mediterranean Sea to the Aegean Sea by strong currents toward the south-east (Corsini & Lefkaditou 1994). A similar distribution trend associated with the sea currents was proposed for O. tuberculata in the Adriatic Sea (Tutman et al. 2008). This species enters the Adriatic, probably from the Mediterranean Sea, following a north-westward current toward the northern Adriatic Sea." (Kim et al. 2017) (Kim et al. 2017; Goncalves 1991; Caballero-Alfonso et al. 2009) 182.6 "Lifespan is estimated at half a year, with spawning occurring throughout the year, with some seasonal variability in activity (Jereb et al. 2010)." (Downes et al. 2017) (Downes et al. 2017) 182.6 "Lifespan is estimated at half a year, with spawning occurring throughout the year, with some seasonal variability in activity (Jereb et al. 2010)." (Downes et al. 2017) (Downes et al. 2017) NA 0 2 1 1 "Males have been observed travelling inside hollow jellyfish (salps)." (Norman and Reid 2000 – Book, p81) Norman and Reid 2000 – Book, p81 1 (Bello 2012) 7 "Ocythoe are reported to feed on pteropod and heteropod molluscs, sardines, and crustaceans" (Jereb et al., 2014) Jereb et al 2014
Tutman et al 2008
Katsanevakis et al 2008		 0 0 4 13 "A considerable number of epipelagic predators reportedly prey on O. tuberculata
including the lancetfishes, Alepisaurus borealis (by Berry, 1955) and A. ferox
(by Rees & Maul, 1956); the tunas, Thunnus alalunga (by Iverson, 1971), T.
thunnus (by Pinkas, 1971), and Germon germon (by Bouxin & Legendre, 1936);
and Risso's dolphin, Grampus griseus (by Joubin, 1900)" (Roper &
Sweeney 1975)		 (Tejerina, 2019) (Blanco
et al., 2006) (Jereb
et al., 2014) (Markaida
& Sosa-Nishizaki, 2010) (Antonelis
et al. 1987) (Downes
et al. 2017) (Battaglia
et al. 2013) (Cardoso
1991) (Lipinski
& David 1990) (Roper
& Sweeney 1975) (Norman
and Reid 2000)		 6 0 not gregarious (inferred from photo and video material) 1 0 1 1 "the only known ovoviviparous cephalopod genus that incubates eggs in the oviducts until paralarvae hatch" (Laptikhovsky & Salman, 2002) (Laptikhovsky & Salman, 2002) 0 "the only known ovoviviparous cephalopod genus that incubates eggs in the oviducts until paralarvae hatch" (Laptikhovsky & Salman, 2002) (Laptikhovsky & Salman, 2002) 31 18 264 101.108 93.058 44 42 NY-W-W
Onychoteuthis banksii Onychoteuthis banksii 200 "Female Ocythoe typically occupy near-surface waters, having been encountered in the upper 10m and captured in plankton hauls and on hook and line. While male Ocythoe have been collected in pelagic tows at the sea surface, they are not restricted to surface waters, having been collected in closing nets at 100 to 200 m." (Jereb et al., 2014) (Laptikhovsky & Salman, 2002; Tejerina, 2019; Jereb et al., 2014; Quetglas et al. 2013; Norman and Reid 2000 – Book, p81) 0 "surface-living species (Nesis 1987)" (Laptikhovsky & Salman, 2002) (Laptikhovsky & Salman, 2002; Tejerina, 2019; Jereb et al., 2014; Quetglas et al. 2013; Norman and Reid 2000 – Book, p81) 1 "Oceanic, mesopelag- ic." (Gomes-Pereira et al., 2016) Gomes-Pereira et al., 2016 40 62 62 N to 22 N (Jereb & Roper, 2010) (Jereb & Roper, 2010) 22 62 N to 22 N (Jereb & Roper, 2010) (Jereb & Roper, 2010) 4 1 “The site of greatest abundance occurs in the Tyrrhenian Sea (station 3) in summer and shifts eastward to the Ionian Sea (station 4) in winter.” (Roper 1974) (Roper 1974) 0 "There is no strong evidence of ontogenic shift as this species grows" (Judkins & VEcchione, 2020) (Judkins & VEcchione, 2020) 1 “O. banksi clearly has a broad vertical range, particularly during the daytime (summer), but heaviest concentrations occur in the upper 100 m both day and night. The range is more restricted at night when nearly the entire population occurs at 25-100 m. As strong, active swimmers the larger juveniles and the adults are able to make occasional excursions to depths of 600-800 m during the day, apparently returning toward the surface waters at night. Winter captures, less numerous than those of summer, occur over a more restricted range of depths (0 – 375 m).” (Roper 1974) (Roper 1974; Watanabe et al., 2006) 365.3 "The lifespan of both sexes is approximately 1 year, but males mature at smaller sizes (i.e. about 250 mm), and younger ages than females, where mantle length at maturity varies between 300 and 350 mm." (Jereb & Roper, 2010) (Arkhipkin 1996a; Arkhipkin and Nigmatullin, 1997)" (Arkhipkin, 2004:345; Jereb & Roper, 2010) 365.3 "The lifespan of both sexes is approximately 1 year, but males mature at smaller sizes (i.e. about 250 mm), and younger ages than females, where mantle length at maturity varies between 300 and 350 mm." (Jereb & Roper, 2010) (Arkhipkin 1996a; Arkhipkin and Nigmatullin, 1997)" (Arkhipkin, 2004:345; Jereb & Roper, 2010) 365.3 "The life span of both sexes is approximately 1 year, but males mature at smaller sizes (i.e. about 250 mm), and younger ages than females, where mantle length at maturity varies between 300 and 350 mm." (Jereb & Roper, 2010) (Nixon & Young, 2003:179; Jereb & Roper, 2010) 200 Sexual maturity: "At 200 days of age, males become functionally mature" (Nixon & Young, 2003:179) (Nixon & Young, 2003:179; Jereb & Roper, 2010) NA 0 3 2 1 "mesopelagic cephalopods such as Japetella heathi and Onychoteuthis banksii appear capable of a dynamic strategy of going transparent when downwelling light is present (to avoid detection from predators looking upwards to detect silhoulettes) and switching to red and black chromatophore pigmentation when no downwelling light is present (to avoid detection by predators using bioluminescent searchlights)." (Hanlon & Messenger, 2018, p. 109) Hanlon & Messenger, 2018, p. 109 1 Present in the genus (Jereb & Roper, 2010) (Jereb & Roper, 2010) 1 "This schooling species frequently is observed “flying” above the surface to escape predators in pursuit" (Jereb & Roper, 2010) Jereb & Roper, 2010 4 "Prey consists of fishes and squids." (Jereb & Roper, 2010) Jereb & Roper 2010
Quetglas et al. 2013		 0 1 Arkhipkin & Nigmatullin (1997) 4 21 "Predators
of this species include: epipelagic fishes, e.g. Pacific pomfret, swordfish,
salmon shark; marine mammals, e.g. dolphins, sperm whales, Guadalupe fur seals
and, most significantly in the central North Pacific Ocean, northern fur seals"
(Jereb & Roper, 2010)		 (Blanco
et al., 2006) (Jereb
& Roper, 2010) (Jereb
& Roper, 2010) (Jereb
& Roper, 2010) (Markaida
& Sosa-Nishizaki, 2010) (Kousteni
et al. 2018) (Battaglia
et al. 2013) (Dede
et al 2016) (Ibanez
et al 2004) (Blanco
et al 1995) (Tsuchiya and Sawadaishi 1997)		 4 3 "schooling species" (Jereb & Roper, 2010) Jereb & Roper 2010 1 "schooling species" (Jereb & Roper, 2010) Jereb & Roper 2010 NA 0 NA 34 27 40.11 38.964 15.8 39 36 68 W-W-NY
Pickfordiateuthis pulchella Pickfordiateuthis pulchella 4000 "occurs from the surface to 150 m depth, but it may be found as deep as 4 000 m." (Jereb & Roper, 2010) (Cairns, 1976; Quetglas et al., 2000; Jereb & Roper, 2010; Sifner et al., 2005; Judkins & Vecchione, 2020; Quetglas et al. 2013; Roper 1974; Bolstad 2007; Zylinski and Johnsen 2011; Vecchione & Roper, 1991, p. 436) 0 "O. banksii has been captured at the surface in all three regions of the Straits and in all three water masses." (Cairns, 1976) [Location: Straits of Florida] (Cairns, 1976; Quetglas et al., 2000; Jereb & Roper, 2010; Sifner et al., 2005; Judkins & Vecchione, 2020; Quetglas et al. 2013; Roper 1974; Bolstad 2007; Zylinski and Johnsen 2011; Vecchione & Roper, 1991, p. 436) 2 "Shallow tropical waters on patch reefs and seagrass beds" (Jereb & Roper, 2010) Jereb & Roper, 2010
Voss 1953		 52 28 28 N to 23 S (Jereb & Roper, 2010) (Jereb & Roper, 2010) -24 "Pickfordiateuthis pulchella analyzed herein were
collected in the northern part of the coast of São Paulo State, southeastern Brazil (ranging from 23°26′S, 44°47′W to 23°55′S, 45°35′W)" (Anderson and Marian 2020)		 (Anderson and Marian 2020) 1 NA 0 NA 0 0 0 3 "It is quite common amongst sea grass as schools of 30-60 individuals occur in close formation, 1-3 m above the bottom [Hochberg and Couch 1971)" (Nixon & Young 2003) Hochberg and Couch 1971 cited in Nixon & Young 2003 1 "It is quite common amongst sea grass as schools of 30-60 individuals occur in close formation, 1-3 m above the bottom [Hochberg and Couch 1971)" (Nixon & Young 2003) Hochberg and Couch 1971 cited in Nixon & Young 2003 NA 0 NA 9 5 9.2 21 NY
Pteroctopus tetracirrhus Pteroctopus tetracirrhus 20 0-20 m (Haimovici et al., 1989) (Haimovici et al., 1989; Arango an Diaz 1996) 0 0-20 m (Haimovici et al., 1989) (Haimovici et al., 1989; Arango an Diaz 1996) 2 "P. tetracirrhus has been captured from 26 m (Voss, 1956) to 677-1097 m (Voss, 1955) on mud and shell bottoms. The GERDA specimen was captured between 357-370 m on a fine mud bottom. " (Cairns, 1976) Cairns, 1976
Haimovici et al., 1989
Jereb et al., 2014		 49 45 45.5 N to 1 S (Jereb et al., 2014) (Jereb et al., 2014) -4 recorded off Canary Islands Tenerife. “inhabiting temperate and tropical waters of the Atlantic Ocean including the Mediterranean Sea. In the Western Atlantic, this species extends from North CaroliNAto the coasts off Uruguay, approximately between 40N° and 4°S. In the Eastern Atlantic it is found from the Azores to the south of the Iberian Peninsula and from the West African coast to the equator, including Madeira and Cape Verde. This species can be found in the Mediterranean Sea…Azores” (Escánez et al. 2018) (Escánez et al. 2018) 3 1095.8 "Lifespan
considered to be 2 or 3 years." (Jereb et al., 2014)		 (Mangold 1965; Quetglas et al. 2009; Jereb et al., 2014) 365.3 1 year (Quetglas et al. 2009) [TB: this seems like the best estimate; the authors are quite confident about a life cycle of 1 year and terminal spawning] (Mangold 1965; Quetglas et al. 2009; Jereb et al., 2014) 730.5 Males approx. 18 months and females approx. 18-24 months with Mangold-Wirz
(1963) as reference in (Nixon, 1969)		 Mangold-Wirz
(1963) as reference in (Nixon, 1969)		 547.9 Males approx. 18 months and females approx. 18-24 months with Mangold-Wirz
(1963) as reference in (Nixon, 1969)		 Mangold-Wirz
(1963) as reference in (Nixon, 1969)		 NA 0 NA 0 1 "Ink sac present." (Jereb et al., 2014) (Jereb et al., 2014) 0 0 2 found in stomachs of bluefin tuNAsouthern Tyrrhenian Sea (Battaglia et al. 2013) (Battaglia et al. 2013) (Goren et al., 2006) 2 0 not gregarious (inferred from photo and video material) NA 0 NA "all current knowledge originated from the Catalan Sea (West Mediterranean). P. tetracirrhus reproduces there in summer" (Laptikhovsky et al. 2014) 22 15 283.528 217.892 21.6 92 70 W-W-NY
Pterygioteuthis giardi Pterygioteuthis giardi 1097 "P. tetracirrhus has been captured from 26 m (Voss, 1956) to 677-1097 m (Voss, 1955) on mud and shell bottoms. The GERDA specimen was captured between 357-370 m on a fine mud bottom. " (Cairns, 1976) (Cairns, 1976; Haimovici et al., 1989; Quetglas et al., 2000; Sifner et al., 2005; Jereb et al., 2014; Laptikhovsky et al. 2014) 25 25-720 m (Haimovici et al., 1989). "There were few small-sized octopuses (<7 cm ML) in the samples, which might indicate that these individuals inhabit rocky grounds that are not accessible to trawlers or waters deeper than the maximum depth sampled (800 m). The species occurred more frequently around the Balearic Islands than along the Iberian Peninsula as they appeared in 20% and 7%, respectively, of the hauls in these areas. The octopus inhabits the lower continental shelf and upper slope in both areas, primarily between 200 and 500 m depth." (Quetglas et al. 2009) (Cairns, 1976; Haimovici et al., 1989; Quetglas et al., 2000; Sifner et al., 2005; Jereb et al., 2014; Laptikhovsky et al. 2014) 1 "oceanic species" (Jereb & Roper, 2010) Jereb & Roper, 2010 95 45 45 N to 40 S (Jereb & Roper, 2010) (Jereb & Roper, 2010) -50 [the Falkland Islands] "The Atlantic range of P. giardi is extensive; the species occurs from 408N to 348S (M. Roeleveld, personal communication), from North American to Europe (e.g. Lu & Clarke, 1975; Roper & Young, 1975), south along the African (e.g. Nesis, 1987) and South American continents, at least to the Falkland Islands (Clarke, 1966)." (Lindgren 2010) (Lindgren 2010) 2 0 "Neither of these species appear to undergo an
ontogenic shift during development (Figures 9A,B) within the
limits of our methods." (Judkins & Vecchione, 2020)		 (Judkins & Vecchione, 2020) 0 "Neither of these species appear to undergo an
ontogenic shift during development (Figures 9A,B) within the
limits of our methods." (Judkins & Vecchione, 2020)		 (Judkins & Vecchione, 2020) 1 "It is mesopelagic and ascends into the
epipelagic zone at night." (Jereb & Roper, 2010)		 (Jereb & Roper, 2010) 1 1 1 flashing pairs of photophores “A brilliant flash could become offensive and so allow a predator to disrupt the prey’s defence and enable the predator to make a successful strike. It was suspected ‘that in the dark waters of the open ocean, preypredator strategies involving luminescent flashes are extremely complex.’ (R.E. Young er al. 1982).” (Nixon & Young 2003) Nixon & Young 2003 2 1 1 "We interpret the standard response as an attempt by the squid to startle or distract a predator momentarily, thus allowing the squid to move quickly beyond the predator's strike distance. The flash may also leave a temporary residual image on predator's visual receptors, making it more difficult to follow the prey's movements. The brilliant flash from one of these squid can produce a temporary blind spot in the eye of the dark-adapted human observer in the region of the retiNAwhere the light is focused." (Young et al 1982) Young et al 1982
Nixon & Young 2003		 1 Present in the family (Jereb & Roper, 2010) (Jereb & Roper, 2010) 0 0 3 "The species is preyed upon by large dolphins
(e.g. Tursiops truncatus) and pelagic fishes." (Jereb & Roper, 2010)		 (Jereb & Roper, 2010) (Markaida et al 2015) 3 0 not gregarious (inferred from photo and video material) NA 0 NA 14 9 5.6 23 NY
Pyroteuthis margaritifera Pyroteuthis margaritifera 2200 “BS 668–2200m” (Urbano and Hendrickx, 2018) [benthic sledge] (Cairns, 1976; Jereb & Roper, 2010; Young et al 1987; Urbano and Hendrickx, 2018) 0 "Roper, Gibbs and Aron (1970) reported a day and night range of 300-35J-500 m and 0-102-250 m respectively for 41 specimens captured with closing nets off Bermuda. The GERDA specimens had a day depth range of 256-297-375 m, a twilight range of 70-759-365 m, and a night range of 45-756-389 m (Fig. 2). " (Cairns, 1976; Jereb & Roper, 2010) (Cairns, 1976; Jereb & Roper, 2010; Young et al 1987; Urbano and Hendrickx, 2018) 1 "is found throughout the water column in the North Atlantic and the Gulf, vertically migrating from the upper mesopelagic zone (~600 m depth) during the day to the surface and epipelagic zone at night (Young and Mangold, 2009)" (Timm et al., 2020) Timm et al., 2020 130 70 70 N to 60 S (Jereb & Roper, 2010) (Jereb & Roper, 2010) -60 70 N to 60 S (Jereb & Roper, 2010) (Jereb & Roper, 2010) 2 1 “In winter, no specimens of P. margaritifera were captured at stations 2 and 3, but the species did occur in about equal abundance at stations 4 and 5. P. margaritifera appears to exhibit an eastward shift of the population in winter, leaving the summertime center of abundance in the Tyrrhenian Sea (station 3) entirely devoid of specimens in winter.” (Roper 1974) (Roper 1974; Roper & Young 1975) 1 Canary Basin, NE Atlantic "undergo diel vertical migrations with almost complete separation of the night (mostly 50-100 to 200 m) and day (mostly 400 to 800 m) habitat layers and ontogenetic vertical migrations. The range of diel migration is 300 – 400 m for juveniles and 400 – 500 m for adults. Ontogenetic descent is manifested in the increasing, during the growth, of the average habituation depth primarily during night time." (Nesis 1993) (Nesis 1993; Roper & Young 1975) 1 " The upper mesopelagic zone is the daytime adult habitat; adults then undergo diel vertical migration into epipelagic waters at night. For example, off Bermuda, closing-net studies showed that P. margaritifera occurs principally at 375 to 500 m during the day and ascends to 75 to 175 m at night. Open net studies off the Canary Islands placed animals at 400 to 800 m during the day and at 50 to 100 m to 200 m at night" (Jereb & Roper, 2010) (Jereb & Roper, 2010; Timm et al., 2020; Cairns, 1976; Nesis 1993) NA 0 2 1 1 "Some photophores are concerned in counter-illumination. Pyroteuthis can respond to changes of intensity of overhead illumination, by adjusting the intensity of its own luminescence, so that its outline is not visible when viewed from below [R.E. Young and Roper 1977). The function of the photophores was examined in enoploteuthids and in Pyroteuthis addolux. The latter adopts an oblique position when illuminated from above, and on increasing the light the squid adjusts its own luminescence so that its outline is invisible when viewed from below. This squid could match the light intensities corresponding to the depth range it inhabits during the day (R.E. Young and Roper 1976). The dorsal and ventral photosensitive vesicles of the extraocular light organs are well sited to ensure a match between downwelling light and that emitted by the animal." (Nixon & Young 2003) Nixon & Young 2003 1 Present in the family (Jereb & Roper, 2010) (Jereb & Roper, 2010) 4 Bear Seamount "feeding primarily on copepods, euphausiids, other small invertebrates, and to a lesser extent small fishes and cephalopods (Passarella & Hopkins 1991)…For the early to mid-life stages represented in our evaluations, these squids appear to be primarily tracking and consuming lower trophic level prey (e.g., zooplankton) as they migrate vertically in the water column" (Staudinger et al. 2019) Staudinger et al 2019
Passarella and Hopkins (1991		 0 0 3 3 made
up about 5% of cephalopod species found in stomachs of demersal sharks
Scyliorhinus canicula and Squalus blainville caught in Aegean Sea (Kousteni et
al. 2018)		 (Kousteni
et al. 2018) (Beasley
et al. 2013)		 2 0 not gregarious (inferred from photo and video material) NA 0 NA 17 10 48.888 43.848 19.4 40 32 24 W-W-NY
Rossia macrosoma Rossia macrosoma 1500 "One hundred and six Pyroteuthis margaritifera (8–33 mm ML)
were distributed throughout the water column (0–1500 m)
both day and night but overall they are nyctoepipelagic
synchronous migrators, with the majority of the population
found in the upper mesopelagic zone during daytime and
moving to the epipelagic zone nightly (Figure 4). Smaller
individuals were living at depth whereas larger individuals were
shallower" (Judkins & Vecchione, 2020)		 (Salman et al., 2003; Cairns, 1976; Jereb & Roper, 2010; Timm et al., 2020; Judkins & Vecchione, 2020; Galil & Goren 1994) 0 "One hundred and six Pyroteuthis margaritifera (8–33 mm ML)
were distributed throughout the water column (0–1500 m)
both day and night but overall they are nyctoepipelagic
synchronous migrators, with the majority of the population
found in the upper mesopelagic zone during daytime and
moving to the epipelagic zone nightly (Figure 4). Smaller
individuals were living at depth whereas larger individuals were
shallower" (Judkins & Vecchione, 2020)		 (Salman et al., 2003; Cairns, 1976; Jereb & Roper, 2010; Timm et al., 2020; Judkins & Vecchione, 2020; Galil & Goren 1994) 2 "Sandy and muddy substrate. Demersal" (Jereb & Roper, 2005) Jereb & Roper, 2005 58 70 70 N to 12 N (Jereb & Roper, 2005) (Jereb & Roper, 2005) (Xavier et al., 2018) (Katsanevakis et al 2008) 12 70 N to 12 N (Jereb & Roper, 2005) (Jereb & Roper, 2005) 4 1 "In the western Mediterranean Rossia macrosoma carries out seasonal migrations between deeper offshore waters in winter and shallower coastal zones for the rest of the year. This migration is partitioned by size such that the largest individuals arrive first in spring, followed by smaller animals in summer." (Jereb & Roper, 2005) (Jereb & Roper, 2005) 1 "In the western Mediterranean Rossia macrosoma carries out seasonal migrations between deeper offshore waters in winter and shallower coastal zones for the rest of the year. This migration is partitioned by size such that the largest individuals arrive first in spring, followed by smaller animals in summer." (Jereb & Roper, 2005) (Jereb & Roper, 2005) 365.3 "Females grow larger than males, and longevity is about 12 months." (Jereb & Roper, 2005) Jereb & Roper (2005) 365.3 "Females grow larger than males, and longevity is about 12 months." (Jereb & Roper, 2005) Jereb & Roper (2005) 334.81294 "Mature males, aged 7 to 8 months carry 85 to 100 spermatophores; females, 8 to 11 months, have about 120 to 150 eggs in their ovaries." (Jereb & Roper, 2005) (Jereb & Roper, 2005) 213.06278 "Mature males, aged 7 to 8 months carry 85 to 100 spermatophores; females, 8 to 11 months, have about 120 to 150 eggs in their ovaries." (Jereb & Roper, 2005) (Jereb & Roper, 2005) NA 0 NA 0 1 Present (Jereb & Roper, 2005) (Jereb & Roper, 2005) 3 Portuguese coast “Rossia macrosoma is an active carnivore, typically catching crustaceans, fish and mollusc prey (Mangold-Wirz 1963) and, therefore, most of its dietary carbon comes from these rich protein diets.” (Rosa et al. 2006) Rosa et al 2006
Xavier et al 2018
Katsanevakis et al 2008		 0 0 3 7 “recorded as prey to a wide variety of predators, among which rank marine mammals (Fiscus 1993; Santos & Haimovici 2001) and commercial (Daly et al. 2001; Velasco et al. 2001; Cabral & Murta 2002) and non-commercial fish species and even crustacea (Bello & Pipitone 2002).” (Rosa et al. 2006) (Rosa
et al. 2006) (Kousteni
et al. 2018) (Bello
1997) (Santos
et al. 2006)		 4 2 when placed in tank with sand, sitting on wall "They were often forming groups of about 5 to 10 animals" (von Boletzky 1973) von Boletzky, S., & von Boletzky, M. V. (1973). Observations sur le développement embryonnaire et postembryonnaire de Rossia macrosoma (Mollusca, Cephalopoda). Helgoländer wissenschaftliche Meeresuntersuchungen, 25, 135-161. 0 when placed in tank with sand, sitting on wall "They were often forming groups of about 5 to 10 animals" (von Boletzky 1973) von Boletzky, S., & von Boletzky, M. V. (1973). Observations sur le développement embryonnaire et postembryonnaire de Rossia macrosoma (Mollusca, Cephalopoda). Helgoländer wissenschaftliche Meeresuntersuchungen, 25, 135-161. NA 0 NA 52 26 192.808 128.436 77 51 W-W
Sandalops melancholicus Sandalops melancholicus 900 "It has been reported at depths ranging between 32 and 900 m, caught more frequently on upper slopes. It has been found in the Aegean Sea and the Ionian Sea, at depths between 72 and 700 m. Mature individuals of both sexes are distributed mainly on upper slopes whereas younger individuals are mainly found on the lower shelf (LEFKADITOU & KASPIRIS, 2004; ROSA et al., 2006)." (Katsanevakis et al 2008) Katsanevakis et al 2008 32 "It has been reported at depths ranging between 32 and 900 m, caught more frequently on upper slopes. It has been found in the Aegean Sea and the Ionian Sea, at depths between 72 and 700 m. Mature individuals of both sexes are distributed mainly on upper slopes whereas younger individuals are mainly found on the lower shelf (LEFKADITOU & KASPIRIS, 2004; ROSA et al., 2006)." (Katsanevakis et al 2008) Katsanevakis et al 2008 1 "encompasses the water column from 0 and 1500 m" (Judkins & Vecchione, 2020) Judkins & Vecchione, 2020 86 44 44 N to 42 S (Jereb & Roper, 2010) (Jereb & Roper, 2010) -42 44 N to 42 S (Jereb & Roper, 2010) (Jereb & Roper, 2010) 3 1 "lives in epipelagic, mesopelagic and bathypelagic zones, following the
general cranchiid pattern of ontogenetic descent. By full growth, animals have descended into the bathypelagic zone beyond
2 000 m depth, where maturation and mating occur." (Jereb & Roper, 2010)		 (Jereb & Roper, 2010; Shea et al. 2017) NA 0 NA 0 1 (Young, 1975) 0 0 1 juvenile beaks found in petrels New Zealand waters (Imber 1978) (Imber 1978) 1 0 not gregarious (inferred from photo and video material) NA 0 NA 7 6 34.2 112 NY
Scaeurgus unicirrhus Scaeurgus unicirrhus 2000 "lives in epipelagic, mesopelagic and bathypelagic zones, following the
general cranchiid pattern of ontogenetic descent. By full growth, animals have descended into the bathypelagic zone beyond
2 000 m depth, where maturation and mating occur." (Jereb & Roper, 2010)		 (Jereb & Roper, 2010; Judkins & Vecchione, 2020; Imber 1978) 0 "encompasses the water column from 0 and 1500 m" (Judkins & Vecchione, 2020) (Jereb & Roper, 2010; Judkins & Vecchione, 2020; Imber 1978) 2 Benthic, sand and coraline bottons (Haimovici et al., 1989) Haimovici et al., 1989 74 45 45 N to 29 S (Jereb et al., 2014) (Jereb et al., 2014) -29 45 N to 29 S (Jereb et al., 2014) (Jereb et al., 2014) 4 1 "Dispersal of species throughout the Atlantic Ocean (according to Briggs 1967) was predominantly from west to east, and this pecies probably therefore originated from the tropical regions of the wester side. S. unicirrhus, as it is now known, has a
planktonic larval stage. Although the duration of this early phase of its life has still not been determined (von Boletzky 1984), it is likely that the species originated in western Atlantic tropical zones, was transported east by surface currents, and colonized the Valdivia Bank. The environment at the Bank is suitable for its survival there." (Sanchez & Alvarez 1988)		 (Sanchez & Alvarez 1988) 730.5 collected two cohorts from southern Tyrrhenian Sea in spring, one of juveniles and other of sub/adults "This fact suggests a two-year life cycle: spawning in spring and early summer, egg development for 2.5 months, paralarvae living comparatively long in the plankton, settlement of juveniles in autumn-winter, and growth for one year or more to reach sexual maturity and reproduce." (Bello 2007) (Bello 2007) 730.5 collected two cohorts from southern Tyrrhenian Sea in spring, one of juveniles and other of sub/adults "This fact suggests a two-year life cycle: spawning in spring and early summer, egg development for 2.5 months, paralarvae living comparatively long in the plankton, settlement of juveniles in autumn-winter, and growth for one year or more to reach sexual maturity and reproduce." (Bello 2007) (Bello 2007) 365.3 Bello (2007) suggests a two-year lifespan with sexual maturity in second year "In spring juveniles range in size from 20 to 42 mm ML. Hence they need to live for one more year to reach sexual maturity and reproduce in the following spring" Bello (2007) 365.3 Bello (2007) suggests a two-year lifespan with sexual maturity in second year "In spring juveniles range in size from 20 to 42 mm ML. Hence they need to live for one more year to reach sexual maturity and reproduce in the following spring" Bello (2007) NA 0 NA 0 1 (Jereb et al., 2014; Bello, 2004) 3 "It feeds on molluscs, crustaceans, and small fishes." (Jereb et al., 2014) (Jereb et al., 2014) 0 0 3 3 made
up about 2 and 17% of cephalopod species found in stomachs of demersal sharks
Scyliorhinus canicula and Squalus blainville caught in Aegean Sea (Kousteni et
al. 2018)		 (Kousteni
et al. 2018) (Di
Lorenzo et al., 2020)		 1 0 not gregarious (inferred from photo and video material) 1 0 1 0 "The egg brooding period, according to Boletzky's (1984) data, may be inferred to last at least 2.5 months at 13-14°C in the Mediterranean." (Bello 2007) (Bello 2007) 1 "The egg brooding period, according to Boletzky's (1984) data, may be inferred to last at least 2.5 months at 13-14°C in the Mediterranean." (Bello 2007) (Bello 2007) 34 27 138.6 136.25 110.25 62 50 NY-W-W
Sepia bandensis Sepia bandensis 800 " inhabited depths from 100 to 800 m." (Quetglas et al., 2000) (Cairns, 1976; Haimovici et al., 1989; Quetglas et al., 2000; Sifner et al., 2005; Silva et al., 2011; Jereb et al., 2014; Laptikhovsky et al. 2014; Giordiano et al. 2010; Katsanevakis et al 2008; Bello 2004) 30 "It is widespread in Hellenic seas, presenting a wider depth range in the Ionian Sea (43-850 m) in comparison to that (30-550 m) in the Aegean Sea, where it seems to be relatively more abundant (Table 3)." (Katsanevakis et al 2008) (Cairns, 1976; Haimovici et al., 1989; Quetglas et al., 2000; Sifner et al., 2005; Silva et al., 2011; Jereb et al., 2014; Laptikhovsky et al. 2014; Giordiano et al. 2010; Katsanevakis et al 2008; Bello 2004) 2 " Coastal shallow waters. Found over coral reefs and sand." (Jereb & Roper, 2005) (Jereb & Roper, 2005
Thompson, 2017		 30 19 19 N to 11 S (Jereb & Roper, 2005) (Jereb & Roper, 2005) -11 19 N to 11 S (Jereb & Roper, 2005) (Jereb & Roper, 2005) 1 273.9 "After 2 to 3 months of continual growth, male dwarf cuttlefish begin displaying sexually dimorphic behaviors (Figure 1B), reaching sexual maturity at around 4 months. Following a period of mating and egg-laying from 4 to 6 months of age, dwarf cuttlefish begin
showing hallmarks of aging, including a reduction in eye-sight and movement, resulting in a failure to catch prey. After a lifetime of continuous growth, dwarf cuttlefish
die at around 9 months of age" (Montague et al. 2021)		 (Montague et al. 2021) 273.9 "After 2 to 3 months of continual growth, male dwarf cuttlefish begin displaying sexually dimorphic behaviors (Figure 1B), reaching sexual maturity at around 4 months. Following a period of mating and egg-laying from 4 to 6 months of age, dwarf cuttlefish begin
showing hallmarks of aging, including a reduction in eye-sight and movement, resulting in a failure to catch prey. After a lifetime of continuous growth, dwarf cuttlefish
die at around 9 months of age" (Montague et al. 2021)		 (Montague et al. 2021) 121.75 "After 2 to 3 months of continual growth, male dwarf cuttlefish begin displaying sexually dimorphic behaviors (Figure 1B), reaching sexual maturity at around 4 months. Following a period of mating and egg-laying from 4 to 6 months of age, dwarf cuttlefish begin
showing hallmarks of aging, including a reduction in eye-sight and movement, resulting in a failure to catch prey. After a lifetime of continuous growth, dwarf cuttlefish
die at around 9 months of age" (Montague et al. 2021)		 (Montague et al. 2021; Montague et al., 2020) 121.8 "Dwarf cuttlefish, Sepia bandensis, are native to the Indo-Pacific, where
tropical water temperatures enable embryos to hatch
after 1 month of development, and animals can reach
sexual maturity in 4 months" (Montague et al., 2020)		 (Montague et al. 2021; Montague et al., 2020) 2 2 1 "It forages over sand, and amongst algae, coral rubble and live coral." (Norman and Reid 2000 – Book, p24) Norman and Reid 2000 – Book, p24 1 "Cuttlefish have a distinct way of hunting, different from their cephalopod cousins. They possess 8 arms and an additional 2 tentacles, which reside in small pockets near the mouth. Similar to a chameleon’s tongue, they extend these two tentacles at high speeds, hitting their target prey item and pulling it into their waiting arms. Feedings are three times daily, and each cuttlefish gets 2-3 shrimp per feeding" (Barber & Ober, 2020) Barber & Ober, 2020 3 3 1 "This species tends to seek cover in crevices rather than jetting away from potential predators." (Norman and Reid 2000 – Book, p24) Norman and Reid 2000 – Book, p24 1 "Colour patterns range from whitc through mottled patterns to uniform dark chocolate brown. The latter colour is used when it hides against the sides of black sea cucumbers." (Norman and Reid 2000 – Book, p24) Norman and Reid 2000 – Book, p24 1 "Dwarf cuttlefish create complex skin patterns for camouflage and social communication. Dwarf cuttlefish can generate a large array of body patterns using variations in color, the spatial frequency of patterns and three-dimensional texture. Cuttlefish skin displays can be associated with camouflage or social behaviors" (Montague et al. 2021) Montague et al. 2021
Hanlon & Messenger, 2018, p. 131		 1 "The S. bandensis diet primarily consists of two species of shrimp: mysid shrimp (Americamysis bahia) for our larval cuttlefish and grass shrimp (Palaemonetes pugio) for our juveniles and adults. " (Barber & Ober, 2020) Barber & Ober, 2020 0 0 1 2 "Multiple adult cuttlefish (or younger animals) can be co-housed, provided the tank is large enough (max. 5 adults per 10 gal), and there are sufficient hiding places (see Methods). By contrast, octopus species are often asocial and cannibalistic." (Montague et al. 2020); "Cuttlefish such as S. bandensis form small aggregations, and hatchlings do well in group housing (Fiorito et al., 2015)." (Bowers et al. 2020); "Dwarf cuttlefish create complex skin patterns for camouflage and social communication. Dwarf cuttlefish can generate a large array of body patterns using variations in color, the spatial frequency of patterns and three-dimensional texture. Cuttlefish skin displays can be associated with camouflage or social behaviors" (Montague et al. 2021) Montague et al. 2020, 2021; Bowers et al. 2020 0 "Multiple adult cuttlefish (or younger animals) can be co-housed, provided the tank is large enough (max. 5 adults per 10 gal), and there are sufficient hiding places (see Methods). By contrast, octopus species are often asocial and cannibalistic." (Montague et al. 2020); "Cuttlefish such as S. bandensis form small aggregations, and hatchlings do well in group housing (Fiorito et al., 2015)." (Bowers et al. 2020); "Dwarf cuttlefish create complex skin patterns for camouflage and social communication. Dwarf cuttlefish can generate a large array of body patterns using variations in color, the spatial frequency of patterns and three-dimensional texture. Cuttlefish skin displays can be associated with camouflage or social behaviors" (Montague et al. 2021) Montague et al. 2020, 2021; Bowers et al. 2020 3 3 1 "We found that, when presented with inaccessible prey, cuttlefish selectively inhibit tentacle strikes without reducing the amount of time oriented towards prey, or total distance moved." Bowers et al. 2020 1 "we showed that juvenile dwarf cuttlefsh demonstrate STM [short term memory] and LTM [long term memory] of the IP [inaccessible prey] experiment by reducing tentacle strikes across two distinct timescales." (Bowers et al. 2021) Bowers et al. 2021 1 "we showed that juvenile dwarf cuttlefsh demonstrate STM [short term memory] and LTM [long term memory] of the IP [inaccessible prey] experiment by reducing tentacle strikes across two distinct timescales." (Bowers et al. 2021) Bowers et al. 2021 NA 6 8 312.100 60 M
Sepia elegans Sepia elegans 10 " Coastal shallow waters. Found over coral reefs and sand." (Jereb & Roper, 2005) (Jereb & Roper, 2005
Thompson, 2017		 0 " Coastal shallow waters. Found over coral reefs and sand." (Jereb & Roper, 2005) (Jereb & Roper, 2005
Thompson, 2017		 2 " living on sandy and sandy – muddy bottoms " (Pierce et al., 2010) Pierce et al., 2010
Jereb et al., 2015
Guerra & Castro 1989		 81 60 From 60 N to 15 S. (Jereb & Roper, 2005) (Jereb & Roper, 2005) -21 "extends through the eastern Atlantic from western Scotland in the north 
to  Namibia  (21S)  in  the  south  and  is  present  throughout  the  Mediterranean  Sea 
(Jereb and Roper, 2005)" (Pierce et al., 2010) but see ""Floating cuttlebones may enter the eastern side of the North Sea and be found stranded
on the Belgian and Dutch coasts (Adam, 1933); however, live S. elegans have never been
found in the North Sea (e.g. Adam, 1933; Roper and Sweeney, 1981; Nesis, 1982/87).
Recent observations confirm this (J. Goud, pers. comm.). The species has been reported
from the Agulhas Bank (37°12’S 22°30’E) by Filippova et al. (1995), but as noted by Roe-
leveld (1998), the position given in Filippova’s paper is incorrect because it lies south
of the Agulhas Bank, in ca. 4000 m depth. " (Jereb et al., 2015)		 (Pierce et al., 2010) (Jereb & Roper, 2005) 1 1 "A spring–summer migration of the whole population to coastal spawning grounds (40–
70 m depth) has been described for the species in the western Mediterranean (Man-
gold-Wirz, 1963a; Guerra, 1992), and similar displacements have been observed in the
Tyrrhenian Sea (off the Tuscany coast; Belcari, 1999a). However, this migratory pattern
does not seem to be displayed in other areas, such as the Ría de Vigo (northwestern
Spain; Guerra and Castro, 1989) or the Sicilian Channel (Jereb and Ragonese, 1991a),
and both presence and absence of migration have been reported for the Adriatic Sea
(Casali et al., 1988; Ciavaglia and Manfredi, 2009). " (Jereb et al., 2015)
 		 (Jereb et al., 2015) (Sánchez et al. 1998) 1 Yes: "It spends the winter in deep water (200–400 m), then migrates into the shallows in spring and summer to spawn." (Jereb & Roper, 2005) Jereb & Roper, 2005) 578.3125 "Lifespan  ranges  between  12  and  19 months." (Pierce et al., 2010) (Pierce et al., 2010; Jereb & Roper, 2005; Jereb et al., 2015) 365.25 "Lifespan  ranges  between  12  and  19 months." (Pierce et al., 2010) (Pierce et al., 2010; Jereb & Roper, 2005; Jereb et al., 2015) 365.25 "Maturity is attained at about 1 year of age." (Jereb & Roper, 2005) (Pierce et al., 2010; Jereb & Roper, 2005) 365.25 "Maturity is attained at about 1 year of age." (Jereb & Roper, 2005) (Pierce et al., 2010; Jereb & Roper, 2005) NA 0 NA 0 33 "It feeds mainly on molluscs, small crustaceans, fishes and polychaetes." (Jereb & Roper, 2005) Jereb and Roper 2005
Guerra 1985
Lelli et al. 2005
Jereb et al 2015		 0 1 Guerra 1985 4 11 Predator species: Common cuttlefish (Sepia officinalis) , European squid (Loligo vulgaris) , Lesser spotted dogfish (Scyliorhinus ca-
nicula) , Bull ray (Pteromylaeus bovinus)
Marbled electric ray (Torpedo mar-
morata) , Thornback ray (Raja clavata) , Common dolphinfish (CoryphaeNAhip-
purus) , European hake (Merluccius merluccius), Greater amberjack (Seriola dumerili) , John Dory (Zeus faber) , Bottlenose dolphin (Tursiops truncatus) (Jereb et al., 2015)		 (Jereb et al., 2015) (Pierce
et al., 2010)		 4 0 not gregarious (inferred from photo and video material) 1 0 1 1 "spawning and nursery areas …are thought to occur farther offshore…minimally influenced by coastal processes" (Dance et al. 2014) Also<"Les Alcyonium palmatum des eaux des environs de Banyuls (entre Collioure et Cerbère) constituent un substrat régulier des pontes de Sepia elegans." [Alcyonium palmatum from the waters around Banyuls (between Collioure and Cerbère) constitute a regular substrate for the spawning of Sepia elegans.] (Bouligand 1962) (Dance et al. 2014) (Bouligand 1962) 79 40 128.7 115.632 55 49 W-W
Sepia officinalis Sepia officinalis 600 "It is common in Hellenic seas where it has been reported at depths of between 20 and 600 m." (Katsanevakis et al 2008) Katsanevakis et al 2008 20 "It is common in Hellenic seas where it has been reported at depths of between 20 and 600 m." (Katsanevakis et al 2008) Katsanevakis et al 2008 2 "It is a benthic, shallow-water species occurring in sea grass beds, empty shells and hard substrates…"(Elsevier, 2014). Elsevier, 2014 46 62 About 62 N to 16 N (Jereb & Roper, 2005) (Jereb & Roper, 2005) 16 About 62 N to 16 N (Jereb & Roper, 2005) (Jereb & Roper, 2005) 1 1 "In the English Channel, S. officinalis hatch inshore at the
beginning of summer (Figure 1). After spending the entire
summer feeding inshore, they migrate offshore to overwinter
in the central western English Channel (Boucaud-Camou &
Boismery, 1991; Dunn, 1999a; Royer et al., 2006) where sea
bottom temperature is favourable (Wang et al., 2003). The following summer is also spent feeding inshore. At the end of this second summer, male specimens finish their maturation (in approximately September). In October, they perform their
second offshore migration to the wintering grounds where
female specimens complete their maturation. The final migration is performed at approximately 20 months old to reach the inshore spawning zones." (Gras et al. 2016)		 (Bettoso et al. 2016; Gras et al. 2016; Dance et al. 2014; Bloor et al. 2013; Wang et al. 2003; Adam, 1937; Pierce et al., 2010; Iglesias et al., 2014; Jereb et al., 2015) 1 Yes: "The species undergoes seasonal migrations between inshore waters during spring and summer and medium shelf grounds (about 100 m depth) during autumn and winter. For example, in the Mediterranean, large individuals leave deeper water early in spring to migrate to shallower water, with males preceding females. This group is followed by a succession of smaller animals throughout the summer. In autumn, gradual decent to deeper water begins." (Jereb & Roper, 2005) (Jereb & Roper, 2005; Alsonso-Fernandez et al. 2021; Ruby and Knudsen 1972; Adam, 1937; Pierce et al., 2010; Jereb et al., 2015; Boyle, 1983) 1 "S. officinalis makes seasonal onshore–offshore migrations (Guerra 2006), likely due to winter
cooling of the waters. Cuttlefish, with their cuttlebone-based buoyancy mechanism,
can also perform daily vertical migration—upward at night for food and downward
in the daytime (Webber et al. 2000)" (Iglesias et al., 2014).		 (Iglesias et al., 2014) 1095.8 "The species lives three years at least" (Ruby and Knudsen 1972) (Jereb & Roper, 2005; Schnell et al., 2021; Le Pabic ete al. 2019; Lin et al. 2019; Wolfram et al. 2006; Zielinski and Poertner 2000; Boyle, 1983; Hanlon & Messenger, 2018, p. 157; Boyle & Rodhouse, 2005; Dunn 1999; Le Goff & Daguzan 1991; Ruby and Knudsen 1972; Pierce et al., 2010; Jereb et al., 2015) 334.8 "By rearing cuttlefish Richard (1971) showed that there is antagonism between
initial growth rate and maximum attained size (or longevity). This also appears
in the natural environment. Here cuttlefish which grow very quickly become group
I breeders, live on average 11 months and die at a maximum size of 19 cm,
whereas those which have a lower initial growth rate live about 22 months and
reach maximum sizes greater than 25 cm for females and 30 cm for males." (Le Goff & Daguzan 1991)		 (Jereb & Roper, 2005; Schnell et al., 2021; Le Pabic ete al. 2019; Lin et al. 2019; Wolfram et al. 2006; Zielinski and Poertner 2000; Boyle, 1983; Hanlon & Messenger, 2018, p. 157; Boyle & Rodhouse, 2005; Dunn 1999; Le Goff & Daguzan 1991; Ruby and Knudsen 1972; Pierce et al., 2010; Jereb et al., 2015) 730.5 "Maturity indices show that cuttlefish are able to
mature before they are 1 year old in the Mediterranean Sea as noted by
Boletzky 1983, while they mature at an age 1 and 2 years in the Atlantic (Gauvrit et al. 1997)." (Guven & Ozbas 2007) [But the real source below]		 (Jereb et al., 2015; Gauvrit et al. 1997; Guven & Ozbas 2007) [But the real source below]; Jereb et al., 2015; Koueta et al. 1995; Koueta et al. 1992) 304.4 "The Sepia officinalis L. used in this study were trawled off the Normandy coast (Bay of
Seine) from July to November. They were in their second year of life, at which time
sexual maturation occurs (Medhioub & Boucaud-Camou, 1989). The males, which were
aged 10-13 months, were undergoing sexual maturation; the females, aged 14-15
months, were in the early previtellogenesis stage" (Koueta et al. 1995)		 (Jereb et al., 2015; Gauvrit et al. 1997; Guven & Ozbas 2007) [But the real source below]; Jereb et al., 2015; Koueta et al. 1995; Koueta et al. 1992) 10 9 1 Ambush: "Prey is often ambushed when these cuttlefishes are partially buried and hidden in sand." (Jereb & Roper, 2005) Jereb & Roper, 2005
(Boal et al. 2000a)." (Boal 2011)		 1 "The loliginid squid the cuttlefishes Sepia officinalis wave their dorsal arm pairs from side to side before attacking prey, which may indicate hypnotizing or luring." (Hoving et al 2013) Hoving et al 2013
Jereb & Roper, 2005
Cooke & Tonkins 2015		 1 "A recent study on the foraging behaviour of common cuttlefish demonstrated that they adapt quickly to changes in their environment using previous experience. Specifically, cuttlefish dynamically change their foraging strategies in response to changes in prey availability and proximate-future expectations (Billard, Clayton, & Jozet-Alves, 2020a)." (Schnell et al., 2020) Schnell et al., 2020 1 "the final (seizure) phase of the cuttlefish [S. officinalis] attack on prawns is also ballistic [i.e., pursuit] (Messenger, 1968). Pursuit may also entail interception of the prey's flight path: this has never been demonstrated reliably in a cephalopod, although cuttlefish [S. officinalis] may anticipate the flight path of a fish by attacking from the front (Messenger, 1968; Neill & Cullen, 1974). Cuttlefish will pursue prey that has moved out of sight" (Hanlon & Messenger, 2018, p. 83) Hanlon & Messenger, 2018, p. 83
Messenger, 1968
(Boal et al. 2000a)." (Boal 2011)
Hanlon & Messenger, 2018, p. 83		 1 "when cuttlefsh (Sepia ofcinalis) are hunting, they often adopt a ‘passing cloud’ display, which involves a white body pattern with dark horizontal bands moving quickly from the mantle towards the arms; this display appears to draw the prey’s attention, causing it to stop moving, making it easier to capture (Hanlon et al. 1988)." (Carreno-Castilla et al 2020) Carreno-Castilla et al 2020
Hanlon & Messenger, 2018, p. 82
Hanlon & Messenger, 2018, p. 81
Cooke & Tonkins 2015
Adamo et al. 2006
Hanlon & Messenger, 2018, p. 81		 1 (2) the “tentacle strike”, used on rapidly escaping preys like shrimps and fish, involves the ejection of the prehensile tentacles, which are rapidly shot out towards the prey and then retracted to bring the prey to the arms and the mouth.” (Zoratto et al. 2018) (Zoratto et al. 2018)
(Messenger 1968; Duval et al. 1984)
(Messenger, 1977)." (O'Brien et al. 2017)
Messenger 1968; Duval et al. 1984)" (Zoratto et al. 2018)
(Duval et al. 1984)” (Agin et al. 2006)		 1 “Seizure”, the final component, can be accomplished by two distinct methods of attack (Messenger 1968; Duval et al. 1984): (1) the “arm grab”, mainly used on crabs (depending on crab size and other factors), involves the use of all the arms without the tentacles once the animal has jumped upon its prey" (Zoratto et al. 2018) (Zoratto et al. 2018)
Nixon & Mangold 1998
MESSENGER, 1968
(Boletzky 1972
Hanlon & Messenger, 2018, p. 82
Messenger 1968; Duval et al. 1984)" (Zoratto et al. 2018)
(Duval et al. 1984)” (Agin et al. 2006)		 1 "In the former [tentacle strike], the tentacles are rapidly extended from a pouch below the eyes toward the prey. The suckers on the tentacle clubs adhere to the prey and bring it to the mouth when the tentacles are retracted. In the jumping attack, the cuttlefish positions itself behind the crab (away from the claws) and pounces on it with all eight arms. It then rotates the crab into a position which allows it to bite the junction between the periopods and the main carapace (Chichery & Chichery, 1988). Their saliva contains a toxin which quickly paralyzes the crab, enabling easy consumption" (O’Brien et al. 2017) O’Brien et al. 2017
Guerra 2006
Agin et al. 2006
Chichery & Chichery 1988		 1 "The tentacles of S. officinalis can reach prey in less than 15 ms. Prey handling is rapid, and neurotoxins secreted by the posterior salivary glands paralyse the prey within 10 s of capture (Hanlon and Messenger, 1996). External digestion does not appear to take place, and (when feeding on crustaceans) many pieces of exoskeleton are ingested (Guerra et al., 1988). " (Jereb et al., 2015) Jereb et al., 2015
Boal, Wittenberg, & Hanlon, 2000; Dickel, 1997)." (O’Brien et al. 2017)
(Boal et al. 2000a)." (Boal 2011)
(Duval et al. 1984)” (Agin et al. 2006)		 1 “In Experiment 2, both cuttlefish from the CREC and from the MBL lowered their consumption of crabs during the day when shrimp were available the following night, while cuttlefish maintained their consumption of crabs during the day when no shrimp were available the following night…Effect sizes for conditions and interactions were greatly above 1 (from 10 to 21 times higher), indicating that cuttlefish alter their foraging behaviour in response to the availability of shrimps the following night, and that this behavioural alteration was even more pronounced across training… Our results could be explained in terms of positive and negative anticipatory contrasts [24]. Indeed, when cuttlefish know that they will not receive any shrimp at night, they would show a positive anticipatory contrast by eating the crabs during the day in anticipation of the absence of a later reward, but when cuttlefish know that shrimp will be distributed at night, they show a negative anticipatory contrast by refraining from eating the crabs, in anticipation of receiving a later reward. This pattern suggests that cuttlefish have rapid and flexible transient foraging strategies in response to changing environmental conditions, previous experience and potentially causal knowledge.” (Billard et al. 2020b) Billard et al. 2020b
Adamo et al. 2006		 10 9 1 "From hatchlings to adults, S. officinalis exhibits a light-induced burying behaviour; most individuals spend the daytime hidden in sand." (Jereb & Roper, 2005) Jereb & Roper, 2005
Di Poi et al., 2013)" (Chabenat et al. 2019		 1 "Ferguson et al. (1994) investigated the CSR of S. offcinalis in more detail and suggested that visual input played a minor role in eliciting the re£ex in cuttlefish. Cuttlefish spend the majority of their time on the sea bottom and their camou£age is largely concerned with matching the diverse backgrounds on which they are found." (Mathger, 2003) Mathger, 2003 1 "Although they did not perfectly match substrate intensity, cuttlefish body pattern intensity matching is likely an important component of an effective camouflage body pattern. In fact, several authors have suggested that intensity matching may be an effective camouflage tactic at greater water depths (Akkaynak et al., 2013; Cott, 1940; Denton and Nicol, 1966; Johnsen, 2001; Lythgoe, 1979; Mäthger et al., 2008) since underwater daylight becomes more and more restricted to the blue–green spectrum with increasing depth (Jerlov, 1976; Tyler and Smith, 1970). Under these blue–green light conditions, color vision may not be as useful for a predator searching for camouflaged prey. In that case, matching the general intensity of the environment may be a successful camouflage tactic." (Buresch et al. 2015) Buresch et al. 2015
Barbosa et al. 2012
Chiao et al. 2011
Allen et al. 2010
Hanlon & Messenger, 2018, p. 112
Ulmer et al. 2013		 1 lab: "Rotation (roll or pitch) of a cuttlefish away from its normal orientation produces countershading reflexes (CSRs) that consist of chromatophore expansion on the ventral body surface…The major role of the CSR in the roll plane is almost certainly that of camouflage from predators. Because of the stability provided by the gasfilled cuttlebone, it is only under unusual circumstances that Sepia become disorientated in the roll plane, and at such times they are probably fleeing rather than attacking. In contrast, the countershading produced in response to rotation in the pitch plane is likely to help avoid detection by both predators and prey, since Sepia often swims up or down through the water column to attack prey (Messenger, 1968)." (Ferguson et al. 1994) Ferguson et al. 1994 1 "In laboratory experiments on cuttlefish, Buresch et al. (Reference Buresch, Mäthger and Allen2011) showed that S. officinalis prefer masquerade of 3D objects over substrate background matching when visual cues on nearby 3D objects are high-contrast and the surrounding substrate is low-contrast (see Fig. 3.16). However, field observations and images suggest that other factors (e.g. number, size, texture and shape of nearby objects) are involved in this visual decision-making process." (Hanlon & Messenger, 2018, p. 113) Hanlon & Messenger, 2018, p. 113
(Packard & Sanders, 1969; Wells, 1978; as cited in Hanlon & Messenger, 2018, p. 127)
Hanlon & Messenger, 2018, p. 129
Ulmer et al. 2013
Hanlon & Messenger, 2018, p. 113		 1 When hunting under lab conditions and presented with an overhead model bird: “Cuttlefish appeared to perceive the sudden appearance of the model bird as a potential threat. All 10 cuttlefish expressed at least some components of the Deimatic Display during the “bird” trials. The Deimatic Display is thought to be an anti-predator response (Hanlon and Messenger, 1996). Therefore, cuttlefish appear to suppress the conspicuous Passing Cloud display (Packard and Hochberg, 1977) in the presence of a potential predator.” (Adamo et al. 2006) Adamo et al. 2006
Hanlon & Messenger, 2018, p. 125
Cartron et al. 2013		 1 analysis of free amino acids in inks of multiple cephs including S. officinalis “Phagomimicry requires that the secretions contain chemicals in absolute and relative concentrations sufficient to mimic the food of predators…Our chemical analysis supports the hypothesis that inking molluscs have the potential to use sensory disruption and/or phagomimicry as a chemical defense” (Derby et al. 2007) Derby et al. 2007
Hanlon & Messenger, 2018, p. 123, 129		 1 "Inking is another form of secondary defence, but in aquaria adults are more likely to emit their ink as a dense cloud, obliterating visibility in all directions for more than a metre, whereas hatchlings and juveniles usually eject small discrete blobs of ink-plus-mucus that stand in the water and attract the predator’s attention to a false cuttlefish while the real cuttlefish escapes. Of course adults can also eject blobs of ink: for a fuller description of inking behaviour see Boycott (1958)" (Hanlon & Messenger 1988) (Adam, 1937; Nixon & Mangold, 1998; Boycott, 1958; Hanlon & Messenger, 1988) 1 Jetting can follow the Deimatic Display if the stimulus (or predator) continues to approach (Hanlon and Messenger, 1996). Every jetting cycle is started by hyperinflation (Packard and Trueman, 1974), and if cuttlefish filled their mantles in advance, they could jet immediately when chromatic displays failed to deter a predator.” (King & Adamo 2006) King & Adamo 2006
(Wilson et al. 2018		 1 "Juvenile cuttlefish can rapidly regenerate amputated arms in a stepwise process" (Callaghan et al., 2019) Callaghan et al., 2019 1 Despite the tendency of animals in Group PVC showing different responses relative to animals in other groups, there were no statistical differences in the secondary defensive behaviors (Fig. S6). It is well known that S. officinalis show different secondary defensive behaviors depending on the predator species (Langridge et al., 2007; Staudinger et al., 2013). However, the European common cuttlefish did not show jet-away/inking behaviors when facing the sea bass in the present study. This observation suggests that the sea bass used in the present study may not be a predator that presents a high predatory risk to this species of cuttlefish. Unlike the puffer fish in Experiment 1, which showed a higher predatory risk to S. pharaonis, the sea bass is likely to be less threatening to S. officinalis embryos in nature. {lee et al 2020) lee et al 2020
Wilson et al. 2018
Langridge 2009
Staudinger et al. 2013
Hanlon & Messenger, 2018, p. 133		 67 "Main crustacean prey items are mysids,
shrimps, prawns, and crabs, but S. officinalis also
feeds upon amphipods, isopods, and ostracods. The
most important bony fishes found in the diet of the
species were gobies, sand eels, whiting and
wrasses, but cuttlefish can also prey upon some
flatfishes. Among the cephalopods main food items
include various sepiolids and sepiids species.
Large cuttlefish are also cannibals, capturing and
eating smaller individuals…S. officinalis shows a wide range
of diets and should therefore be considered as a
trophic opportunist" (Guerra 2006)		 Jereb and Roper 2005
Schnell et al., 2021
Mzaki et al 2017
Guerra 2006
Pinczon du Sel & Daguzan 1997
Guerra 1985		 0 1 Mzaki et al. 2017
Pinczon du Sel & Daguzan 1997
Guerra 1985		 7 49 (Jereb et al., 2015) Predators: Black-mouthed dogfish (Galeus
melastomus) , Blainville's dogfish (Squalus
blainville) , Bluespotted seabream (Pagrus
caeruleostictus) , Blue shark (Prionace glauca) , Bull ray (Pteromylaeus bovineus) , Lesser spotted dogfish (Scyliorhi-
nus canicula) , Pelagic stingray (Pteroplatytry-
gon violacae) , Smooth-hound (Mustelus mus-
telus), Ballan wrasse (Labrus bergylta) , Bib (Trisopterus luscus) , Black seabream (Spondylio-
soma cantharus) , Black-bellied angler (Lophius
budegassa), Brill (Scophthalmus rhombus), Comber (Serranus cabrilla), Common pandora (Pagellus
erythrinus), Conger eel (Conger conger), Dusky grouper (Epinephelus
marginatus), European barracuda (Sphy-
raeNAsphyraena), European hake (Merluccius
merluccius), European seabass (Dicentrar-
chus labrax), Fourspot megrim (Lepidorhom-
bus boscii), Greater amberjack (Seriola
dumerili) , Longfin gurnard (Aspitrigla ob-
scurus) , Megrim (Lepidorhombus whiffi-
agonis), Monkfish (Lophius piscatorius), Pollack (Pollachius pollachius) , Red gurnard (Aspitrigla cuculus), Silver-cheeked toadfish (Lago-
cephalus sceleratus) , Spanish bream (Pagellus
acarne) , Spotted flounder (Citharus lin-
guatula) , Swordfish (Xiphias gladius), Turbot (Scophthalmus maximus), Twaite shad (Alosa fallax), Yellow-mouth barracuda (Sphy-
raeNAviridensis), Yellow-stripe barracuda (Sphy-
raeNAchrysotaenia) , Atlantic grey seal (Halichoerus
grypus), Monk seal (Monachus mona-
chus), Bottlenose dolphin (Tursiops
truncatus), Risso’s dolphin (Grampus
griseus), Oceanic striped dolphin (Sten-
ella coeruleoalba) 		(Blanco et al., 2006) (Jereb & Roper, 2005) (Martins et al. 2018)  (Tuncer & Kabasakal 2016) (Gueera &
González 2011) (Guerra 2006) (Blanc & Daguzan 1999) (Salman et al., 2001) (Andaloro
& Pipitone 1997) (Young & Cockcroft 1995) (Kear et al., 1995) (Pierce
et al., 2010) (Jereb et al., 2015) (Hanlon & Messenger, 2018, p. 145)		 4 1 "They appear to be solitary animals: only single individuals were ever observed underwater in their natural habitat. Whenever a group of hatchlings or juveniles from the laboratory was released underwater, its members always moved away in different directions, showing no tendency to school or move in pairs as adults sometimes do. In the laboratory, young Sepia tolerated fairly crowded rearing conditions; sometimes they aggregated beneath an incoming jet of water but usually their distribution in a tank was random. They showed no signs of intraspecific behaviour, in contrast to mature cuttlefish whose remarkable sexual displays are well known." (Hanlon & Messenger 1988). possibly social juveniles "Despite being held in groups in captivity, cuttlefish (Cephalopoda: Sepiidae) have long been considered rather asocial animals. However, reports of breeding aggregations and one recent schooling observation from the wild started to bring this characterisation into question. Following this, we here present 10 observations of the European cuttlefish Sepia officinalis (Linnaeus 1758) forming groups of up to 30 individuals along the South Coast of the UK. The majority of the observed cuttlefish appeared to be juveniles or subadults and showed different shoaling orientations, such as linear or spherical-shaped formations. This indicated the grouping behaviour did not derive from coincidental accumulations. No mating or courtship behaviour could be identified in these groups, and as all observations were made in August or September, and therefore outside their mating season (March to June), it is unlikely that reproductive behaviour motivates these aggregations. As S. officinalis is known to migrate to deeper overwintering grounds in autumn, we propose that cuttlefish may temporarily form groups in late summer/early autumn as part of their migration pattern, and that their shoaling behaviour likely offers similar fitness benefits as in other migrating shoaling species." (Drerup & Cooke 2021) Hanlon & Messenger 1988; Drerup & Cooke 2021 0 "They appear to be solitary animals: only single individuals were ever observed underwater in their natural habitat. Whenever a group of hatchlings or juveniles from the laboratory was released underwater, its members always moved away in different directions, showing no tendency to school or move in pairs as adults sometimes do. In the laboratory, young Sepia tolerated fairly crowded rearing conditions; sometimes they aggregated beneath an incoming jet of water but usually their distribution in a tank was random. They showed no signs of intraspecific behaviour, in contrast to mature cuttlefish whose remarkable sexual displays are well known." (Hanlon & Messenger 1988). possibly social juveniles "Despite being held in groups in captivity, cuttlefish (Cephalopoda: Sepiidae) have long been considered rather asocial animals. However, reports of breeding aggregations and one recent schooling observation from the wild started to bring this characterisation into question. Following this, we here present 10 observations of the European cuttlefish Sepia officinalis (Linnaeus 1758) forming groups of up to 30 individuals along the South Coast of the UK. The majority of the observed cuttlefish appeared to be juveniles or subadults and showed different shoaling orientations, such as linear or spherical-shaped formations. This indicated the grouping behaviour did not derive from coincidental accumulations. No mating or courtship behaviour could be identified in these groups, and as all observations were made in August or September, and therefore outside their mating season (March to June), it is unlikely that reproductive behaviour motivates these aggregations. As S. officinalis is known to migrate to deeper overwintering grounds in autumn, we propose that cuttlefish may temporarily form groups in late summer/early autumn as part of their migration pattern, and that their shoaling behaviour likely offers similar fitness benefits as in other migrating shoaling species." (Drerup & Cooke 2021) Hanlon & Messenger 1988; Drerup & Cooke 2021 23 18 1 “Habituation was noted by a logarithmic decrease in the occurrence of certain responses over the course of 30 exposures (30 min) of repeated 200 Hz tone stimuli. This decrease was notable in the more dramatic escape responses (inking and jetting), and for large body patterning changes…evasion responses suggest that the cuttlefish initially reacted to the stimulus as they would react to a predator or other form of danger, and that sound detection could be a mechanism for predator detection in these animals. After several exposures and no imminent threat, the number of escape responses decreased, suggesting the cuttlefish were able to filter out the ‘irrelevant’ acoustic stimuli, allowing for a refocusing of sensory mechanisms”. However “Total response inhibition was never reached; individuals repeatedly exhibited a ‘stereotyped startle’ response” (Sampson et al. 2014) Sampson et al. 2014 1 "Two groups of cuttlefish (Sepia officinalis) were used to demonstrate classical conditioning in
this species and to determine whether the resulting approach response would be that of sign
tracking or goal tracking. For cuttlefish in the paired condition, a flashing light was presented
at one end of a long tank followed by food dropped into the center of the tank. For cuttlefish in
the unpaired condition, food was dropped into the center of the tank either before or after the
flashing-light stimulus. Paired cuttlefish oriented to the light, positioned themselves within
striking distance, and occasionally attacked the light. Unpaired cuttlefish showed no reliable
response to either stimulus. The results demonstrate that cuttlefish are capable of signal
learning and that, under the conditions tested, cuttlefish sign tracked."		 Purdy et al. 1999 1 "By changing the right/left location of the two visual cues at the end of acquisition, the authors showed that half of the cuttlefish consistently swam in the opposite direction of the rewarded goal compartment; this result indicates that cuttlefish were using visual cues to solve the task. The other cuttlefish consistently swam in the correct direction of the rewarded goal compartment; this result means that these cuttlefish were using a motor sequence to solve the task. These two experiments successfully demonstrated that cuttlefish can use either visual cues or motor sequence to solve the same spatial task." (Dickel et al. 2013) Dickel et al. 2013 1 "both common cuttle fish, Sepia officinalis, and bobtail squid, EuprymNAscolopes, can learn rapidly to inhibit predatory behaviour through aversive associative learning (Agin et al., 1998, 2006a, 2006b; Zepeda, Veline, & Crook, 2017)." (Schnell et al., 2020) Schnell et al., 2020 1 "In experiment 3, one out of four common cuttlefish (S. officinalis) selected the correct escape doorway in 13 out of 15 mixed trials. From these experiments, we conclude that these cephalopods can conditionally discriminate." Hvorecny et al. 2007 1 lab evidence for episodic memory: “during Phase 1, the cuttlefish learn which prey is associated with which visual cue; during Phase 2, the cuttlefish are tested either after a short (1 hour) or a long (3 hour) delay. After the short delay, the non-preferred prey is available but not the preferred prey. After the long delay, both prey are available….it is clear that all three cuttlefish switched their preference to choosing the visual cue associated with non-preferred prey when tested after 1 hour, and continued to choose the visual cue associated with preferred prey after 3 hours (Figure 1D). This pattern of choices demonstrates What-WhereWhen memory — ‘what’ (prey type) were located ‘where’ (location of the visual cue) and ‘when’ (time elapsed since they previously ate)” (Jovet-Alves et al. 2013) Jovet-Alves et al. 2013 1 "Brightness-discrimination was learnt by all subjects within 27 ± 4.58 trials (mean ± s.e.) over 1–4 days and discrimination-reversal was learnt by all subjects within 46 ± 6.70 trials (mean ± s.e.) over 2–6 days. The mean number of trials to reach the learning criterion differed significantly between the two different phases (t 5 = − 5.92, p < 0.01; electronic supplementary material, figure S4). Performance in the brightness-discrimination phase was correlated positively with performance in the discrimination-reversal phase (Pearson’s r = 0.861, p < 0.05; figure 3). Schnell et al. 2021 1 "S. officinalis can extract the vertical and horizontal components of a learned 2D coordinate, and use them independently. Furthermore, we showed that cuttlefish prefer vertical to horizontal spatial information. The ability to recall the vertical and horizontal components of a location independently from each other suggests that cuttlefish encode the two types of information separately…During the conflict tests, seven out of eight animals preferred the visual cue at the correct height to the visual cue at the correct horizontal location…hydrostatic pressure could make navigation in the vertical plane more informative for cuttlefish, as suggested for fish, because it allows for accurate depth detection…Vertical displacement is also associated with a higher predation risk for both cuttlefish and the benthic fish Corydoras aeneus, which surfaces to breathe (Domenici et al., 2007, 2015; Kramer and McClure, 1980). This may contribute to assign a stronger ecological value to information in the vertical dimension. (Davis et al., 2014; Holbrook and Burt de Perera, 2011)" (Scata et al. 2016) Scata et al. 2016 1 “the PT [‘prawn in a tube’] learning procedure is probably largely associative, with the presence of the tube (perceived by both visual and tactile cues) being largely associated with the absence of food reward at each attempted capture. It is of crucial importance to highlight that PT learning in cuttlefish is likely to involve both tactile and visual memory systems of the brain…the inferior frontal lobe should also be considered as a structure of interest. Indeed, this lobe receives tactile proprioceptive inputs and sends axons to the subvertical and superior frontal (probably the anterior part) lobes. One can consider those lobes receiving inputs from the optic lobes as a network of multimodal integration that may integrate visual and tactile information in PT learning.” (Dickel et al 2013) Dickel et al 2013; Mather 1986 1 "Cuttlefish that attack bitter prey immediately show disgust responses: they attempt to remove the quinine from the prey, direct the prey, pulse jets of water towards the prey, and move away from it. Moreover, even by the end of the second trial, experimental cuttlefish gradually increase their attack latency until they stop attacking, generally within 10 trials. This inhibition of predatory behaviour towards the originally preferred prey persists for at least 72 h afterwards, and in a choice test individuals continue to attack a different prey" (Agin et al. 2006) Agin et al. 2006 1 “the PT [‘prawn in a tube’] learning procedure is probably largely associative, with the presence of the tube (perceived by both visual and tactile cues) being largely associated with the absence of food reward at each attempted capture. It is of crucial importance to highlight that PT learning in cuttlefish is likely to involve both tactile and visual memory systems of the brain…the inferior frontal lobe should also be considered as a structure of interest. Indeed, this lobe receives tactile proprioceptive inputs and sends axons to the subvertical and superior frontal (probably the anterior part) lobes. One can consider those lobes receiving inputs from the optic lobes as a network of multimodal integration that may integrate visual and tactile information in PT learning.” (Dickel et al 2013); "Sepia used tactile cues also, because animals hovered off the bottom and extended their arms onto gravel and beads. Their rejection of gravel as a substrate for digging may have been based on tactile learning, which has been proven in Octopus (Wells & Wells, 1957). Tactile cues assisted in terminating the response, because it reoccurred in shallow sand (not covered enough) and occurred in uncovered (dorsal mantle only)." (Mather 1986) Dickel et al 2013; Mather 1986 0 "Male cuttlefish presented with a mirror also reliably showed the Intense Zebra Display (37 of 48 presentations; Adamo & Hanlon, Reference Adamo and Hanlon1996) emphasising the role of visual stimuli as well as the lack of recognition of ‘self’. This display is highly stereotyped and ritualised, and contains at least nine separate signals: two postural, one locomotor, one textural and five chromatic (Box 7.1)." (Hanlon & Messenger, 2018, p. 158) Hanlon & Messenger, 2018, p. 158 1 raised and tested in lab: “We compared animals under three experimental conditions. In two, there was a unique training trial of different duration (5 or 20 min), a third group served as controls. Our results demonstrate that the control situation failed to reduce the level of attack; in contrast the short-term retention, obtained after a single learning trial, is related to a specific short-term memory process…Our data on pooled animals confirm earlier observations with a 24 h retention delay (Messenger, 1973a) in showing that there is indeed an inverse relationship between the number of strikes and the duration of the initial training trial, i.e. an increase in the time of exposure to the glass tube containing the prey corresponding to a proportional decrease in the number of strikes exhibited by animals during the retention phase” (Agin et al. 1998) Agin et al. 1998 1 lab, juvenile cuttlefish: “Our data show that giving a naturally nonpreferred prey item to cuttlefish just once on Day 3 induced a good long-term retention of this novel preference in 7-day-old animals. This would be the first demonstration of a form of long-term memory in cuttlefish during the first week of life”(Darmaillacq et al. 2004a) Darmaillacq et al. 2004a 1 "Imprinting and perceptual learning for prey preference have been demonstrated in S. officinalis. As reported by Wells (1958) and others (e.g., Darmaillacq, Chichery, Poirier, & Dickel, 2004; Darmaillacq, Chichery, & Dickel, 2006; Darmaillacq, Chichery, Shashar, & Dickel, 2006; Guibé et al., 2012) hatchlings have an “innate” preference for shrimp or shrimp-shaped objects (but see below). However, that this preference can be overridden by chemical and/or visual exposure to crabs shortly after hatching…prey preference could also be induced on the basis of brightness contrast: Where naïve cuttlefish preferred dark to white crabs as their initial meal, embryos and hatchlings exposed to white crabs later preferred these over dark crabs (Guibé et al., 2012). This demonstrates that S. officinalis is able to learn about multiple characteristics of prey (shape and/or contrast). Moreover, cuttlefish pre- or postnatally exposed to white crabs preferred black crabs over shrimp, indicating that S. officinalis will generalize the characteristics of a learned preference to the closest alternative if the preferred item is not available (Guibé et al., 2012)" (O'Brien et al. 2017) O'Brien et al. 2017 1 "Cuttlefish delayed gratification when it led to a prey item of higher quality and they were able to maintain delays for periods of up to 50–130 s." (Schnell et al., 2021) Schnell et al. 2021 1 cuttlefish “quickly learned the simple task of exiting a straight alley with escape as the only motivation…Additionally, the lack of effect of maze reversal suggests that the cuttlefish did not rely on visual features around the laboratory to solve this simple maze problem“, learned to escape from vertical wall two-choice maze (Karson et al. 2003) Karson et al. 2003 1 Dickel et al. 2013 1 "Despite unparalleled flexibility in pattern production, S. officinalis usually produced cryptic body patterns with a high degree of bilateral symmetry. The deimatic display, in particular, was often expressed asymmetrically, which is contrary to the predicted use of symmetry in colour patterns involved in predator–prey interactions." Langridge 2005 5 0 "To date, very few interspecific associations (excluding parasitism) have been reported for this species." (Guerra 2006) Guerra 2006 0 "Sepia officinalis are highly visual animals and tolerate high densities in captivity, and this has led previous experimenters to assume that the species is social. However, Boal’s (Reference Boal 1996) multiple experiments showed, among other things, no evidence of recognition of individual mates, and thus cast doubt on the possibility that S. officinalis possesses any complex social recognition." (Hanlon & Messenger 2018); "this species of cuttlefish appears to lack social recognition, including the capability to identify individual mates or rivals (Boal 1996, 2006; Palmer et al. 2006). Instead, sexual recognition is rather primitive. Although there is some evidence that females can recognize other females using visual cues (Palmer et al. 2006), cuttlefish typically identify the gender of their conspecifics using a signal-response system… Male cuttlefish presented with a mirror also reliably showed the intense zebra display (37 of 48 presentations; Adamo and Hanlon 1996), emphasizing the role of visual stimuli as well as the lack of self-recognition." (Allen et al. 2017) Boal 1996 cited in Allen et al. 2018, Hanlon & Messenger 2018 0 lab "Young cuttlefish, up to an age of about 4 months, show no kind of visual intraspecific communication. Mature females when placed together in a confined space generally sit motionless, often close together, showing one of the chronic patterns" (Hanlon & Messenger 1988), but also lab: “We found that females displayed a newly described body pattern termed Splotch toward their mirror image and female conspecifics, but not to males, prey or inanimate objects…providing evidence that a channel for female signaling exists. The ability of female cuttlefish to reliably express the Splotch body pattern when exposed to female conspecifics also suggests that cuttlefish can recognize other females, even when they are presented as mirror images…Females frequently presented Splayed arms to other females and to their mirror image. Splayed arms is typically expressed during foraging behaviour and to potentially threatening stimuli (Hanlon and Messenger 1988). This result suggests females treat other females as potential threats until they receive a specific signal, possibly Splotch, from that conspecific. Mirror images elicit longer and more frequent bouts of Splayed arms, probably because mirror images never withdraw or cease to make the display before the target female does…Although suggestive, the existence of a reliable channel for female-specific signaling is not sufficient to demonstrate that female cuttlefish communicate with one another. For example, the Splotch body pattern could be a reflection of the animal’s internal state (see discussion in Boal et al. 2004) with no effect on other cuttlefish…may serve a similar purpose [deterring attacks or cannibalism when males signal to e/o] between females in S. officinalis.” (Palmer et al. 2006) & “Experiment 1 addressed whether the presence of conspecifics affected the visible, achromatic body patterns of males…No single factor or variable differed significantly between conditions (F5,25 2.37, p 0.05). Thus, we have no strong evidence that cuttlefish body patterns differ depending upon the particular conspecific(s) viewed…Suggestive differences in body patterns were found in response to (1) the sight of an empty tank as compared to other cuttlefish (all variables, Fig. 2), (2) the sight of females as compared to males (zebra patterning, Fig. 2c, f), and (3) the sight of one as compared to two individuals of the same sex (deimatic spots, Fig. 2d).” (Boal et al. 2004) & “[when presented with a film that distorts polarization between test subject and conspecific] We found no evidence that polarized signals affected male behavior. In the first set of trials, males responded primarily to the sex of the other cuttlefish rather than to whether or not polarization was distorted..females were more active when viewing another female through a clear barrier that transmitted normal polarization than through a filter that distorted polarization…females oriented in parallel more often and showed more zebra banding to males viewed through a clear barrier than through a polarization-distorting filter (Fig. 4c). Although none of these data were significant alone, when taken together they suggest that polarization information could be important to female receivers…polarization of body patterns could be a signal to conspecifics that means ‘I am here,’ or ‘stay away'. The sender could transmit this visual signal without breaking crypsis to potential predators.” (Boal et al. 2004) & "They appear to be solitary animals: only single individuals were ever observed underwater in their natural habitat. Whenever a group of hatchlings or juveniles from the laboratory was released underwater, its members always moved away in different directions, showing no tendency to school or move in pairs as adults sometimes do. In the laboratory, young Sepia tolerated fairly crowded rearing conditions; sometimes they aggregated beneath an incoming jet of water but usually their distribution in a tank was random. They showed no signs of intraspecific behaviour, in contrast to mature cuttlefish whose remarkable sexual displays are well known." (Hanlon & Messenger 1988) (Hanlon & Messenger 1988) 1 In Lab Waving 1st or 8th arms "Warning signal by dominant male to subordinates. Frequently leads
to circling/chasing/jetting/inking" (Cooke & Tonkins 2015) & In lab with captive-raised: "In experiment 5, I did not see any reduction in the number of male–male contests across observation periods; indeed, during experiment 2 when males were housed in small groups for several months, they displayed to each other almost continuously. These experiments provided no evidence for discrimination of familiar from unfamiliar individuals…That male cuttlefish form dominance hierarchies (experiment 5) provides evidence for the assertion that dominance hierarchies are not dependent upon individual or even class recognition (Archawaranon et al. 1991; Zayan 1992)…Cuttlefish have adequate visual receptivity to distinguish each other’s banding patterns (Budelmann 1994). Whether they are incapable of perceiving and remembering these distinctions or whether they are simply not motivated to discriminate is unknown.” (Boal 1996) but see "They appear to be solitary animals: only single individuals were ever observed underwater in their natural habitat. Whenever a group of hatchlings or juveniles from the laboratory was released underwater, its members always moved away in different directions, showing no tendency to school or move in pairs as adults sometimes do. In the laboratory, young Sepia tolerated fairly crowded rearing conditions; sometimes they aggregated beneath an incoming jet of water but usually their distribution in a tank was random. They showed no signs of intraspecific behaviour, in contrast to mature cuttlefish whose remarkable sexual displays are well known." (Hanlon & Messenger 1988)		 13 “We formally define a body pattern in terms of the probabilities that various skin features are expressed, and use Bayesian statistical methods to estimate the number of distinct body patterns and their visual characteristics. For the dataset of cuttlefish coloration patterns recorded in our laboratory, this statistical method identifies 12–14 different patterns, a number consistent with the 13 found by Hanlon and Messenger…the expression of cuttlefish body patterns in our dataset does not follow Zipf’s law…One possible reason why cuttlefish signals do not resemble natural languages is that many patterns are used primarily for camouflage. More importantly, with 54 different components, it is potentially possible that each signal could communicate up to 54 bits of information, more than enough for very sophisticated communication. By contrast, rather than 54 bits, because of redundancy, we found an estimate of 3.4 bits per signal…Although small, for a relatively non-social species, this capacity may be all that S. officinalis requires.” (Crook et al. 2002) (Crook et al. 2002) 1 "undergoes a long seasonal migration between deeper waters in
winter and shallower coastal areas in spring, using estuaries as
mating and spawning grounds (Neves et al., 2009)." (Rosa et al. 2015) but see "No known spawning aggregations exist for S. officinalis, and it seems that such observations will be fortuitous because only small groups of this species have ever been
observed, even in spawning season (early spring), along European coastlines and the Mediterranean Sea." (Allen et al. 2017)		 (Rosa et al. 2015) & (Gras et al. 2014) & (Nixon & Mangold 1998) 1 “One female was placed in the tank on Day 1 (mantle length, 13.8cm). 1 day later, 2 males were added…The female died on Day 8 and was replaced with a new female…we did not see any behaviours that we could classify as male courtship, as has been observed in field studies on two related cuttlefish species. Males approached females and attempted to grasp them…in most cases, however, the female blew water at the male (3/12 mating events), jetted away from the male (11/12), or inked (8/12)…The female either created a cloud of ink around her and then slowly swam away (n=3), leaving the males in the ink, or the female would send the ink in one direction and jet in another direction (m=5)…males attempted to remain close (less than 3 mantle lengths away) to the female after copulation and chased away the other male if he approached. The female frequently attempted to get away from the male during this time. Males sometimes exhibited the typical male-male agonistic display, the Intense Zebra display, to the female during these events.” (Adamo et al. 2000) (Adamo et al. 2000) & (Hanlon et al. 1999) & (Boal 1997) 0 “There were no indications of any form of courtship between the female and either male, which also agrees with laboratory observations (Boal 1996, 1997; Boal et al. 1999; Hanlon et al. 1999; Adamo et al. 2000). Moreover, the female did not pay attention to the fights and even wandered away twice; this also seems typical of this species (Boal 1997)… Boal (1997) found that female choice in Sepia officinalis was not based on winners of fights as much as it was on apparent chemical cues: the male that had mated most recently was the choice of females in that laboratory study.” (Allen et al. 2017) & “One female was placed in the tank on Day 1 (mantle length, 13.8cm). 1 day later, 2 males were added…The female died on Day 8 and was replaced with a new female…we did not see any behaviours that we could classify as male courtship, as has been observed in field studies on two related cuttlefish species. Males approached females and attempted to grasp them…in most cases, however, the female blew water at the male (3/12 mating events), jetted away from the male (11/12), or inked (8/12)…The female either created a cloud of ink around her and then slowly swam away (n=3), leaving the males in the ink, or the female would send the ink in one direction and jet in another direction (m=5)…males attempted to remain close (less than 3 mantle lengths away) to the female after copulation and chased away the other male if he approached. The female frequently attempted to get away from the male during this time. Males sometimes exhibited the typical male-male agonistic display, the Intense Zebra display, to the female during these events.” (Adamo et al. 2000) (Allen et al. 2017) & (Adamo et al. 2000) & (Hanlon et al. 1999) 1 "Male cuttlefish compete vigorously for female mates. Larger males win most fights; for example, larger males forced smaller males to retreat in 11 of 14 laboratory trials (Adamo and Hanlon 1996). When sexually mature male cuttlefish fight, key components of the intense zebra display, such as the darkness of one male’s face, can predict whether their behavior will escalate to a violent agonistic encounter (Adamo and Hanlon 1996). Initially, each animal adopts a body pattern composed of light and dark zebra stripes. Next, one or both animals extends his fourth arm (the only sexually dimorphic character) toward the other male. They also produce a dark ring around the eye, sometimes with a unilaterally dilated pupil. When both males maintain a dark face (many brown chromatophores expanded), the encounter is likely to escalate to grappling or biting." (Allen et al. 2017) & "Two male cuttlefish (consort, male 1; intruder, male 2) competing for a female in the Turkish Aegean Sea. A, The consort male approached the female. B, The consort male copulated with the female for about 4 min. C, The consort male remained in proximity to the female for more than 2 min. D, The consort male and the female were approached by male 2, and they began chromatic and postural signaling. The consort male showed a dark face during an intense zebra display (E) and then inked and jetted away (F). G, The original consort male later approached male 2, who was now the consort, and engaged in another agonistic bout. H, Male 1 darts in very closely to male 2 in an attempt to take over consortship once again. I, Attached to each other, the males tossed the trio in several barrel rolls; the female broke loose and jetted toward the surface; and the males rolled a few additional turns before separating and swimming away…the winning male (male 2 in bouts 1 and 2, male 1 in bout 3) performed more medium-level and high-level aggressive behaviors than the losing male (fig. 2A). For the three bouts, the rate of escalation appeared to follow a pattern of successive progression (fig. 2A). Additionally, the zebra stripe contrast measurements showed that once a male signaled strong zebra banding, he usually continued to signal at this level of aggression. This pattern of escalation suggests that male Sepia officinalis use sequential signaling during contests." (Allen et al. 2017) (Allen et al. 2017) 1 "an elaborate and ritualized courtship, which incorporates stereotyped visual displays and ‘mate guarding’" (Jereb & Roper, 2005) & "S. officinalis males present a striped body pattern (intense zebra display) to other
males; larger and darker males are dominant in such interactions (Boal 1997). How-
ever, females show consistent preference not for the larger or darker males, but for
the most recently mated males and those showing fewer zebra displays (Boal 1997).
She suggested that females used chemical cues rather than visual ones in assessing
males. In captivity, males may initiate copulation without obvious courtship (which
females may avoid through escape responses), and male–male aggression occurs.
The extent to which these behaviours are artefacts of spatial constraint in captivity
is not known (Adamo et al. 2000) and may pose challenges for cultivation. Hanlon
et al. (1999a) observed that males initiate mating in the head-to-head posture, and
then direct jets of water on at the female’s buccal membrane, likely to flush sper-
matangia placed there by previous mating, and then transfer their own spermatangia
to the buccal membrane using the hectocotylus. The male then manipulates the
spermatangium on the female to break it open so that sperm are released. Females
appear to terminate mating and are then guarded briefly by the male" (Iglesias et al., 2014).		 (Jereb & Roper, 2005) & (Iglesias et al., 2014). 0 "No parental care has been reported in the species. " (Jereb et al., 2015) (Jereb et al., 2015) & (Lee, 2020) 0 "No parental care has been reported in the species. " (Jereb et al., 2015) (Jereb et al., 2015) & (Lee, 2020) 1392 922 433.236 396.2 144.096 185 80 70 W-NY-W
Sepia orbignyana Sepia orbignyana 242 Caught in depths of 20–242 m with mean depth of 72 m (Silva et al., 2011) (Jereb & Roper, 2005; Salman et al., 1998; Quetglas et al., 2000; Sifner et al., 2005; Silva et al., 2011; Guerra et al. 2016; Xavier et al. 2016; Bloor 2016; Jereb & Roper, 2005; Sanchez & Martin 1993; Ward & Boletzky 1984; Ruby and Knudsen 1972; Boyle, 1983; Jereb et al., 2015; Elsevier, 2014; Moussa et al. 2019; Bettoso et al. 2016; Boyle & Rodhouse, 2005) 0 “Sepia officinalis Linnaeus 1758, the common cuttlefish of the eastern Atlantic and Mediterranean, occurs on sandy to muddy bottoms from the coastline to 200 m, and is most abundant in the upper 100 m…” (Boyle & Rodhouse, 2005) (Jereb & Roper, 2005; Salman et al., 1998; Quetglas et al., 2000; Sifner et al., 2005; Silva et al., 2011; Guerra et al. 2016; Xavier et al. 2016; Bloor 2016; Jereb & Roper, 2005; Sanchez & Martin 1993; Ward & Boletzky 1984; Ruby and Knudsen 1972; Boyle, 1983; Jereb et al., 2015; Elsevier, 2014; Moussa et al. 2019; Bettoso et al. 2016; Boyle & Rodhouse, 2005) 2 "Sepia orbignyaNAis a demersal species that lives mainly on sandy and sandy-muddy bottoms. [..] In the Sea of Marmara the species can occur in brackish waters" (Jereb & Roper, 2005) Jereb & Roper, 2005
Pierce et al., 2010
Jereb et al., 2015		 72 55 About 55 N to 17 S (Jereb & Roper, 2005) (Jereb & Roper, 2005) -17 "The pink cuttlefish, Sepia orbignyaNAFérus-
sac in d’Orbigny, 1826, is found in the
Northeast Atlantic and throughout the Med-
iterranean (Nesis, 1982/1987; Roper et al.,
1984; Guerra, 1992; Reid and Jereb, 2005)
(Figure 8.2), although the northern limits of
its distribution are unclear. It is reported in
the Irish Sea and the English Channel (Nesis,
1982/87, Reid and Jereb, 2005), but it is not
included among the species listed by Massy
(1928) for the Irish coast. In addition, alt-
hough Adam (1952) indicates that its distribution extends to the Arcachon Basin (west-
ern France), there is no mention of its presence along the east coast of England in old
records (e.g. Grimpe, 1925), and the reference to the English Channel by Norman (1890,
p. 484 in Stephen, 1944) apparently is a misquotation. Strandings of cuttlebones of this
species are, however, known from North Sea coasts (e.g. the Netherlands; Cadee, 2002).
The species can be found south along the French and Spanish coasts (Morales, 1958;
Adam and Rees, 1966), in the Bay of Biscay, south to ca. 17°S (southern Angola; Adam,
1962). Sepia orbignyaNAis widely distributed throughout the Mediterranean Sea (Man-
gold and Boletzky, 1987; Bello, 2004; Salman, 2009) including western and central Med-
iterranean parts (Mangold-Wirz, 1963a; Sánchez, 1986a, Belcari and Sartor, 1993; Jereb
and Ragonese, 1994; Giordano and Carbonara, 1999; Relini et al., 2002; Cuccu et al.,
2003a), the Adriatic Sea, although it is only rarely caught in the northern part (Casali et
al., 1998; Krstulović Šifner et al., 2005; Piccinetti et al., 2012), the Ionian Sea (Tursi and
D’Onghia 1992; Lefkaditou et al., 2003a; Krstulović Šifner et al., 2005), the Aegean Sea,
and the Levant Basin (D’Onghia et al., 1992; Salman et al., 1997; 1998; Lefkaditou et al.,
2003b). The species has been recorded also in the Sea of Marmara (Katağan et al., 1993;
Ünsal et al., 1999)." (Jereb et al., 2015)		 (Jereb et al., 2015) (Katsanevakis et al 2008) 1 0 "In the Mediterranean, males and females are usually found together throughout the
year, and no onshore spawning migrations have been reported (Mangold-Wirz, 1963a;
Jereb and Ragonese, 1991a; D’Onghia et al., 1992; Ciavaglia and Manfredi, 2009). " (Jereb et al., 2015)		 (Jereb et al., 2015) 547.875 "Lifespan is estimated to be 12–18 months (Mangold-Wirz, 1963a), although preliminary
estimates from analysis of length-frequency distributions suggested a longer life, i.e.
ca. 3 years (Ragonese and Jereb, 1991). As in many cephalopods, length-frequency dis-
tributions generally are polymodal, although it is difficult to identify microcohorts, and
growth estimation by means of length-frequency methods is generally unreliable for
cephalopods (e.g. Caddy, 1991). " (Jereb et al., 2015)		 (Jereb et al., 2015) 365.25 "Lifespan is estimated to be 12–18 months (Mangold-Wirz, 1963a), although preliminary
estimates from analysis of length-frequency distributions suggested a longer life, i.e.
ca. 3 years (Ragonese and Jereb, 1991). As in many cephalopods, length-frequency dis-
tributions generally are polymodal, although it is difficult to identify microcohorts, and
growth estimation by means of length-frequency methods is generally unreliable for
cephalopods (e.g. Caddy, 1991). " (Jereb et al., 2015)		 (Jereb et al., 2015) 304.375 "Mature males, aged 6 or 7 months, carry about 100 spermatophores; females of 9 or 10 months of age, carry around 400 eggs." (Jereb & Roper, 2005) (Jereb & Roper, 2005) 182.625 "Mature males, aged 6 or 7 months, carry about 100 spermatophores; females of 9 or 10 months of age, carry around 400 eggs." (Jereb & Roper, 2005) (Jereb & Roper, 2005) 1 1 1 "The only information about the mode of prey catching by S. orbignyaNAis provided by Boletzky, who reared some specimens of this species in the aquarium (Boletzky 1988a). He observed that these cuttlefish caught their prey by tentacle ejection only (Boletzky pers. comm.)." (Bello & Piscitelli 2000) Bello & Piscitelli 2000 NA 0 11 "Diet consists mainly of crustaceans, but fish and cephalopods make up a minor component." (Jereb & Roper, 2005) Jereb and Roper 2005
Belivermis et al., 2019
Jereb et al 2015		 0 0 6 4 "Known predators of Sepia orbignyaNAin the Mediterranean Sea and Northeast Atlantic. " Lesser spotted dogfish (Scyliorhinus canicula), Thornback ray (Raja clavata) , Black-bellied anglerfish (Lophius budegassa) , Audouin’s gull (Larus audouinii) (Jereb et al., 2015) (Jereb
et al., 2015) (Pierce
et al., 2010)		 3 0 not gregarious (inferred from photo and video material) NA 0 NA "spawning and nursery areas …are thought to occur farther offshore…minimally influenced by coastal processes" (Dance et al. 2014) 56 22 245.344 207.196 80 62 W-W
Sepia plangon Sepia plangon 550 "from very shallow (15–20 m; Belcari
and Sartor, 1993; Bello et al., 1994; Casali et al., 1998; Ciavaglia and Manfredi, 2009; I.
Sobrino, pers. comm.) down to maximum recorded depths of 565 m in the Mediterra-
nean Sea (Cuccu et al., 2003a) and 580 m in the eastern Atlantic (Gulf of Cádiz, I. So-
brino, pers. comm.). However, the species is most abundant between 50 and 250 m
throughout the Mediterranean Sea, as confirmed by numerous studies (Mangold-Wirz,
1963a; Adam, 1952; Lumare, 1970; Restuccia and Ragonese, 1986; Sánchez, 1986a; Au-
teri et al., 1988; Mannini and Volpi; 1989; Soro and Piccinetti-Manfrin, 1989; Katağan
and Kocatas, 1990; Repetto et al., 1990; Jereb and Ragonese, 1991a; Würtz et al., 1991;
D’Onghia et al., 1992; Belcari and Sartor, 1993; Katağan et al., 1993; Salman et al., 1997;
Quetglas et al., 2000; González and Sánchez, 2002; Ciavaglia and Manfredi, 2009). There is also a major concentration of the species between 340 and 360 m in the Gulf of Cádiz
(eastern Atlantic, I. Sobrino, pers. comm.). As in S. elegans, it is the peculiar structure of
the cuttlebone, which is small, narrow, and with closely packed septa and modified
sutures (Ward, 1991), that allows this species to reach these remarkable depths and to
be among the deepest living Sepia species known. Records from below 450 m are scarce
(e.g. Lefkaditou et al., 2003a), and captures below 550 m extremely so, because that is
the depth below which the shell starts to implode (Ward and Boletzky, 1984). " (Jereb et al., 2015) Since this is unpublished data it is not used: "Its bathymetric range extends from a few meters down to middle slope, with its deepest record from a haul at 700-770 m in the southern Aegean Sea (MEDITS – International bottom trawl survey in the Mediterranean, unpublished data)." (Katsanevakis et al 2008)		 Quetglas et al., 2000
Sifner et al., 2005
Jereb & Roper, 2005
Katsanevakis et al 2008		 15 "from very shallow (15–20 m; Belcari
and Sartor, 1993; Bello et al., 1994; Casali et al., 1998; Ciavaglia and Manfredi, 2009; I.
Sobrino, pers. comm.) down to maximum recorded depths of 565 m in the Mediterra-
nean Sea (Cuccu et al., 2003a) and 580 m in the eastern Atlantic (Gulf of Cádiz, I. So-
brino, pers. comm.). However, the species is most abundant between 50 and 250 m
throughout the Mediterranean Sea, as confirmed by numerous studies (Mangold-Wirz,
1963a; Adam, 1952; Lumare, 1970; Restuccia and Ragonese, 1986; Sánchez, 1986a; Au-
teri et al., 1988; Mannini and Volpi; 1989; Soro and Piccinetti-Manfrin, 1989; Katağan
and Kocatas, 1990; Repetto et al., 1990; Jereb and Ragonese, 1991a; Würtz et al., 1991;
D’Onghia et al., 1992; Belcari and Sartor, 1993; Katağan et al., 1993; Salman et al., 1997;
Quetglas et al., 2000; González and Sánchez, 2002; Ciavaglia and Manfredi, 2009). There is also a major concentration of the species between 340 and 360 m in the Gulf of Cádiz
(eastern Atlantic, I. Sobrino, pers. comm.). As in S. elegans, it is the peculiar structure of
the cuttlebone, which is small, narrow, and with closely packed septa and modified
sutures (Ward, 1991), that allows this species to reach these remarkable depths and to
be among the deepest living Sepia species known. Records from below 450 m are scarce
(e.g. Lefkaditou et al., 2003a), and captures below 550 m extremely so, because that is
the depth below which the shell starts to implode (Ward and Boletzky, 1984). " (Jereb et al., 2015)		 Jereb & Roper, 2005 2 Reef (Chung et al. 2023) Chung et al. 2023 24 -10 10 S to 34 S (Jereb & Roper, 2005) (Jereb & Roper, 2005) -34 10 S to 34 S (Jereb & Roper, 2005) (Jereb & Roper, 2005) 1 304.375 "In captive animals, the life cycle is less than 10 months." (Jereb & Roper, 2005) (Jereb & Roper, 2005) 304.375 "In captive animals, the life cycle is less than 10 months." (Jereb & Roper, 2005) (Jereb & Roper, 2005) NA 0 2 2 1 behavior in drop-chain trawl “Squid that were observed within schools generally maintained the same behaviour as their cohorts while within the bosom of the trawl mouth…Swimming behaviours were similar for both individual squid and squid observed in schools. For both groups, the most squid were observed to “drift”, 60.3 and 63.6%, respectively (Table 3). Similarly for the two groups, the second most frequent swimming behaviour was “jet-swim”, which made up 38.9% of observed individual squid and 36.4% of squid in schools…Many squid did appear already herded at first observation, and all but six squid were orientated with their mantle away from the trawl, as was described by Glass et al. (1999). For the six squid, their orientations appeared to be a reaction to a predator, summer flounder. As the summer flounder moved in front of the trawl, these squid changed their orientation, and some inked—the only squid observed to ink in this study.” (Bayse et al. 2016) Bayse et al. 2016 1 behavior in drop-chain trawl “Squid that were observed within schools generally maintained the same behaviour as their cohorts while within the bosom of the trawl mouth…Swimming behaviours were similar for both individual squid and squid observed in schools. For both groups, the most squid were observed to “drift”, 60.3 and 63.6%, respectively (Table 3). Similarly for the two groups, the second most frequent swimming behaviour was “jet-swim”, which made up 38.9% of observed individual squid and 36.4% of squid in schools…Many squid did appear already herded at first observation, and all but six squid were orientated with their mantle away from the trawl, as was described by Glass et al. (1999). For the six squid, their orientations appeared to be a reaction to a predator, summer flounder. As the summer flounder moved in front of the trawl, these squid changed their orientation, and some inked—the only squid observed to ink in this study.” (Bayse et al. 2016) Bayse et al. 2016 1 NSW, Australia observed preying on White’s seahorse, Hippocampus whitei (Harasti et al. 2014) Harasti et al. 2014 0 0 1 2 "The mourning cuttlefish Sepia plangon Gray, 1849 is unusual among cuttlefish as its individuals associate both during and outside the mating season (McBride, 2005). In S. plangon there is a seasonal shift in both group size and composition, with adults pairing during the breeding season (June–November) and larger groups (n . 6) of immature cuttlefish forming in the post- spawning season (McBride, 2011). Behavioural investigations have shown that the pairs of mature cuttlefish are likely to be mating associations (McBride, 2011); however, the function of the juvenile groupings remains unknown." (Umbers et al., 2016) Umbers et al., 2016 0 "The mourning cuttlefish Sepia plangon Gray, 1849 is unusual among cuttlefish as its individuals associate both during and outside the mating season (McBride, 2005). In S. plangon there is a seasonal shift in both group size and composition, with adults pairing during the breeding season (June–November) and larger groups (n . 6) of immature cuttlefish forming in the post- spawning season (McBride, 2011). Behavioural investigations have shown that the pairs of mature cuttlefish are likely to be mating associations (McBride, 2011); however, the function of the juvenile groupings remains unknown." (Umbers et al., 2016) Umbers et al., 2016 8 2 1 "Deceptive displays, where males simultaneously pro- duced male patterns on one side of the mantle and mimicked the female on the other (figure 1), were pro- duced only by males when courting a single female in the presence of a single rival male, despite the wide range of social groups observed. While the deception was confined to a single social context, it was common within that context, with 39 per cent of groups contain- ing males employing the tactic (figure 2). This result is compelling, because most males were observed either by themselves or in a mating pair; thus, if the behaviour occurred at random, it should have been observed more readily in the most common social contexts." (Brown et al., 2012) Brown et al. 2012 1 Skin display: "mourning cuttlefish, Sepia plangon, can produce a courtship display towards a female on one side of their body while displaying female patterning on the other side of their body towards a rival male, presumably in an attempt to prevent the rival from interfering with their courtship behaviour (Brown, Garwood, & Williamson, 2012)." (Schnell et al., 2020) Schnell et al., 2020 6 1 "While evidence
for non-kin group benefits (Clutton-Brock, 2009) is absent for
this species, there is some evidence of intraspecific communica-
tion in S. plangon (Brown, Garwood & Williamson, 2012). If S.
plangon groups are cooperative, their formation may be driven
by mutualistic mechanisms such as resource dependence or vigi-
lance. " (Umbers et al., 2016)		 (Umbers et al., 2016) 1 "Around Hong Kong, animals migrate to shallower waters during the mating season, where large numbers of adults congregate in 40 to 80 m on the continental shelf from November to February." "During the spawning season, 4 or 5 adults congregate, sometimes near a boulder. " (Jereb & Roper, 2005) (Jereb & Roper, 2005) 1 Skin display: "mourning cuttlefish, Sepia plangon, can produce a courtship display towards a female on one side of their body while displaying female patterning on the other side of their body towards a rival male, presumably in an attempt to prevent the rival from interfering with their courtship behaviour (Brown, Garwood, & Williamson, 2012)." (Schnell et al., 2020) (Schnell et al., 2020) 1 "Populations are male biased and males
compete for receptive females, display mate guarding,
displace rivals and interrupt courtship attempts
(C. McBride & J. E. Williamson, unpublished data)." (Brown et al., 2012)		 (Brown et al., 2012) 1 "Populations are male biased and males
compete for receptive females, display mate guarding,
displace rivals and interrupt courtship attempts
(C. McBride & J. E. Williamson, unpublished data)." (Brown et al., 2012)		 (Brown et al., 2012) 1 "they use dynamic
visual displays for intraspecific communication. Males
generally exhibit a pattern of pulsating stripes on the
mantle during interspecific interactions, whereas
females have characteristic mottled camouflage color-
ation [12]." " This study, however, is
the first account of simultaneous dual gender signalling
in cephalopods combining both female mimicry and
laterally split displays." (Brown et al., 2012)		 (Brown et al., 2012) 9 9 872.5 386.3 455.62 107 72.9 71.1 C20-C20-C20
Sepietta oweniana Sepietta oweniana 83 "(1–83 m)" Chung et al., 2023 Chung et al., 2023
Jereb & Roper, 2005		 1 "(1–83 m)" Chung et al., 2023 Chung et al., 2023
Jereb & Roper, 2005		 2 "It prefers soft, muddy substrates throughout its distributional range, and it is often found on shrimp fishing grounds. Tolerance to salinity variations seems lower than that observed in other bobtail squids and S. oweniaNAhas never been found in brackish waters." (Jereb & Roper, 2005) Jereb & Roper, 2005
Jereb et al., 2015
Boyle, 1983		 59 71 "A single specimen of Sepietta oweniaNAwas captured in the Tromsø Bank (70854′ N 19846.8′ E; depth 175 m) (Figure 1) by a demersal trawl" (Golikov et al. 2014) (Golikov et al. 2014) 12 70 N to 12 N (Jereb & Roper, 2005) (Jereb & Roper, 2005) 1 1 "Animals in wild populations feed mainly upon crustaceans; a specific preference for the euphasiid Maganyctiphanes norvegica in north Atlantic waters and the decapod Pasiphaea sivado in the northern Tyrrhenian Sea has been observed, supporting hypothesized trophic migrations of S. oweniaNAin response to prey abundance and distribution." (Jereb & Roper, 2005) (Jereb & Roper, 2005) 1 "Sepietta oweniaNAis found most of the year on soft bottoms at depths of 50-600 m along the northeast Atlantic. Migration to shallower water for spawning has been reported from the Catalonian Sea and a similar situation is probable in the Skagerrak…" (Boyle, 1983). (Boyle, 1983; Jereb & Roper, 2005; Belcari et al. 2015; Jereb et al., 2015; ) 273.9 "The whole life cycle is, therefore, rather short and may range between 6 and 9 months, depending mainly on the time of embryonic development." (Jereb & Roper, 2005) (Jereb & Roper, 2005; Czudaj et al. 2011; Jereb et al., 2015) 182.6 "The whole life cycle is, therefore, rather short and may range between 6 and 9 months, depending mainly on the time of embryonic development." (Jereb & Roper, 2005) (Jereb & Roper, 2005; Czudaj et al. 2011; Jereb et al., 2015) 182.625 Portuguese waters “Putative age estimates derived from increment counts in our study correspond to an age estimate of 4.5 – 6 months for mature and spent females and 5 – 5.5 months for males with the highest reproductive system weight. This is in agreement with the time-span of 5 – 6 months for the development from hatching to spawning reported from captivity studies in species of the genera Sepietta and Sepiola (Boletzky 1975).” (Czudaj et al. 2011) (Jereb & Roper, 2005; Jereb et al., 2015; Deickert 2009; Czudaj et al. 2011; Jereb et al., 2015) 121.7 " Animals mature at 4–5 months of age" (Jereb et al., 2015) (Jereb & Roper, 2005; Jereb et al., 2015; Deickert 2009; Czudaj et al. 2011; Jereb et al., 2015) 1 1 1 observations from feeding in tank "Adults generally launch their attacks from positions near the bottom. When a buried squid spots prey it emerges from the bottom (Fig. la). The squid leaves the bottom using the fins for forward swimming and keeps the funnel jet pointing downward. The arms form a tight cone around the slightly protruding tentacles (Fig. 1b). When approaching the prey, both juveniles and adults bleach while searching for an optimal position, slightly below the prey's level. Normally the squid has a dense reddish-brown pigmentation with iridescent green areas around the eyes. At an optimal attacking distance, normally about the same as the squid's total body length, the tentacles are thrown forward and simutaneously the red brown pigmentation returns. When the tentacles are shot out, the arms fling open with suckers extended and form an umbrella (Fig. lc). This sequence of movements prepares for the seizure of the prey and counteracts the momentum, created by the tentacular movement" (Bergstrom 1985) Bergstrom 1985 0 “Observations on newly hatched and young S. oweniaNAattacking air bubbles and floating particles suggest that prey capture is intially indiscriminate; as long as the object is of the appropriate size and keeps moving it is attacked. Likely prey organisms in the natural habitat are larval pandalid shrimps, small euphasiids, small mysids, and large copepods” (Bergstrom 1985) Bergstrom 1985 3 3 1 "Feeding occurs primarily from dusk to dawn, with adult animals spending the day buried in the bottom substrate." (Jereb & Roper, 2005) Jereb & Roper, 2005 1 This free-swimming (planktonic-nektobenthic) strategy gradually changed with age and they eventually spent more time on or buried in the shell sand. This burying behaviour is similar to that described for other sepiolids by Boletzky & Boletzky (1970)" (Boyle, 1983). Boyle, 1983 1 observations from feeding in tank "Adults generally launch their attacks from positions near the bottom. When a buried squid spots prey it emerges from the bottom (Fig. la). The squid leaves the bottom using the fins for forward swimming and keeps the funnel jet pointing downward. The arms form a tight cone around the slightly protruding tentacles (Fig. 1b). When approaching the prey, both juveniles and adults bleach while searching for an optimal position, slightly below the prey's level. Normally the squid has a dense reddish-brown pigmentation with iridescent green areas around the eyes. At an optimal attacking distance, normally about the same as the squid's total body length, the tentacles are thrown forward and simutaneously the red brown pigmentation returns. When the tentacles are shot out, the arms fling open with suckers extended and form an umbrella (Fig. lc). This sequence of movements prepares for the seizure of the prey and counteracts the momentum, created by the tentacular movement" (Bergstrom 1985) Bergstrom 1985 9 "Animals in wild populations feed mainly upon crustaceans; a specific preference for the euphasiid Maganyctiphanes norvegica in north Atlantic waters and the decapod Pasiphaea sivado in the northern Tyrrhenian Sea has been observed" (Jereb & Roper, 2005) Jereb and Roper 2005
Xavier et al 2018
Jereb et al 2015		 0 0 6 22 "Giant
red shrimp (Aristaeomorpha foliacea) Bello and Pipitone (2002); Deep-water rose
shrimp (Parapenaeus longirostris) Sobrino et al. (2005); Black-mouthed dogfish
(Galeus melastomus) Macpherson (1981), Kabasakal (2002), Fanelli et al. (2009);
Kitefin shark (Dalatias licha) Matallanas (1982); Lesser spotted dogfish
(Scyliorhinus canicula) Macpherson (1981); Rabbit fish (Chimera monstrosa)
Bello (1997); Smooth lanternshark (Etmopterus pusillus) Xavier et al. (2012);
Thornback ray (Raja clavata) Kabasakal (2002), Šantić et al. (2012); Velvet
belly lanternshark (Etmopterus spinax) Macpherson (1981), Sartor (1993), Neiva
et al. (2006); Atlantic cod (Gadus morhua) Bergström and Summers (1983);
Blue-mouth (Helicolenus dactylopterus) Neves et al. (2011); European hake
(Merluccius merluccius) Carpentieri et al. (2000, 2005); Haddock (Melanogrammus
aeglefinus) Bergström and Summers (1983); Swordfish (Xiphias gladius) Salman
(2004); Cetacea Common dolphin (Delphinus delphis) Santos et al. (2004a, 2013); Harbour porpoise (PhocoeNAphocoena)
Börjesson et al. (2003), Santos et al. (2005b); Harbour seal (Phoca vitulina)
Brown et al. (2001); Striped dolphin (Stenella coeruleoalba) Würtz and Marrale
(1993)” (Belcari et al. 2015)		 (Jereb & Roper, 2005) (Xavier et al., 2018) Sifner &
Vrgoc (2009) (Kousteni et al. 2018) (Bello 1997) (Belcari et al. 2015) (Jereb
et al., 2015) (Boyle, 1983). (Katsanevakis et al 2008)		 5 2 "some evidence of aggregation in the sepiolines…1250 Sepietta oweniaNA[captured] in one trawl lasting just 1h (Belcari er al. 1989)" (Nixon & Young 2003) Nixon & Young 2003 0 "some evidence of aggregation in the sepiolines…1250 Sepietta oweniaNA[captured] in one trawl lasting just 1h (Belcari er al. 1989)" (Nixon & Young 2003) Nixon & Young 2003 2 0 2 1 Portuguese waters “Sepietta oweniaNAhas been noted to associate with Rondeletiola minor, Sepietta neglecta and with Rossia macrosoma in the upper depth strata of the latter’s vertical range (Lefkaditou & Kaspiris 2005; Reid & Jereb 2005) and the same was encountered in the Portuguese coast (Rosa et al. 2006).” (Czudaj et al. 2011) Czudaj et al. 2011 1 "A major spawning location for S. oweniaNAin the Catalan Sea is between 90 to 120 m depth (Deickert & Bello, 2005)." (Deickert 2009) & Sardinian waters “A total of 166 females (ML: 12.90-30.80 mm; TW: 1.29-8.30 g) and 188 males (ML: 13.40-29.10 mm; TW: 1.45-6.33 g) of S. oweniaNAwere found between 53 and 598 m depth, but the bulk of the sample came from 300-500 m in both summer and winter. Mature and maturing specimens were present in both seasons.” (Cuccu et al. 2010) (Deickert 2009 0 "Copulation involves the insertion of the male hectocotolyzed arm into the female‘s mantle cavity to transfer spermatophores. The copulation of sepiolid squid follows the general pattern reported by Racovitza (1894) for Sepiola atlantica d’Orbigny (cited by Mangold-Wirz. 1963) and later amplified by Mangold-Wirz (1963) for Sepiola rondeleti Steenstruup and S. oweniana. Mating is completed in a short period of time and appears to be quite brusque; no initial courtship behaviour has been reported. The male grabs the female head on, turns her upside down and holds her firmly with the lateral arms. The male’s dorsal arms are then put into the female’s mantle cavity. Spermatophores are deposited close to the orifice of the left oviduct where they burst and spermatozoa are stored in the bursa copulatorix. A mated female can often be recognized by the remains of spermatophores in the left side of the mantle cavity. Mangold-Wirz (1963) found that most females with large eggs had already mated when caught (spring and summer)" (Boyle, 1983) (Boyle, 1983) 50 33 72.168 44.268 33 23 W-W
Sepiola affinis Sepiola affinis 1000 "an epipelagic–mesopelagic species, occurring within a wide depth range, i.e. from close to the surface (8 m) down to over 1 000 m. In the North Atlantic it is most common between 50 and 300 m and in the Mediterranean between 100 to 200 and 400 m." (Jereb & Roper, 2005) (Jereb & Roper, 2005; Quetglas et al., 2000; Sifner et al., 2005; Silva et al., 2011; Villanueva 1992; Czudaj et al. 2011; Cuccu et al. 2010; Jereb et al. 1997; Jereb et al., 2015; Boyle, 1983; Katsanevakis et al 2008) 8 "an epipelagic–mesopelagic species, occurring within a wide depth range, i.e. from close to the surface (8 m) down to over 1 000 m. In the North Atlantic it is most common between 50 and 300 m and in the Mediterranean between 100 to 200 and 400 m." (Jereb & Roper, 2005) (Jereb & Roper, 2005; Quetglas et al., 2000; Sifner et al., 2005; Silva et al., 2011; Villanueva 1992; Czudaj et al. 2011; Cuccu et al. 2010; Jereb et al. 1997; Jereb et al., 2015; Boyle, 1983; Katsanevakis et al 2008) 2 "mostly on sandy or sandy-muddy substrates" (Jereb & Roper, 2005) Jereb & Roper, 2005 10 45 45 N to 35 N (Jereb & Roper, 2005) 35 45 N to 35 N (Jereb & Roper, 2005) 1 2 2 1 "Boletzky and Boletzky (1970) described how Sepiola aflinis rise out of the substrate and strike passing prey" (Hanlon & Messenger, 2018, p. 82) Hanlon & Messenger, 2018, p. 82 1 "During prey capture the species exhibits a great variety of colour patterns, with rapid colour changes." (Jereb & Roper, 2005) Jereb & Roper, 2005 3 2 1 "The patterns called A and Al are the most frequently observed during phase 2/3, i.e. during tentacle ejection, when the highest numbers of colour changes are observed. Considering that during this phase the predator's attention must be fully focussed on the prey (binocular fixation), so that its alertness against other predators momentarily decreases, one may wonder whether these rapid colour changes represent an adaptation to minimize the risk for the predator of being attacked by another animal. This could indeed be achieved by rapid changes having a "startling" effect on would-be predator…It indeed blends perfectly with sandy substrates on which the animal is mostregularly exposed to potential predators. Interestingly the "basic pattern" neverappears flash-like. Among the other patterns that have been found to occur inchronic expression, "ALL DARK" might be "cryptic" in certain circumstances,but it certainly appears as "striking" as many complex patterns when seen againsta light background. In contrast to the "basic pattern", "ALL DARK" mayindeed be expressed flash-like. Similarly "ALL LIGHT" tends to be expressedvery briefly. Clearly, they do not appear in the most active phase of prey capture,but when expressed very briefly, "ALL DARK" and "ALL LIGHT" might havea startling effect similar to what has been suggested above for phase 2/3" (Mauris 1989) Mauris 1989 1 Present (Jereb & Roper, 2005) (Jereb & Roper, 2005) 1 "The patterns called A and Al are the most frequently observed during phase 2/3, i.e. during tentacle ejection, when the highest numbers of colour changes are observed. Considering that during this phase the predator's attention must be fully focussed on the prey (binocular fixation), so that its alertness against other predators momentarily decreases, one may wonder whether these rapid colour changes represent an adaptation to minimize the risk for the predator of being attacked by another animal. This could indeed be achieved by rapid changes having a "startling" effect on would-be predator…It indeed blends perfectly with sandy substrates on which the animal is mostregularly exposed to potential predators. Interestingly the "basic pattern" neverappears flash-like. Among the other patterns that have been found to occur inchronic expression, "ALL DARK" might be "cryptic" in certain circumstances,but it certainly appears as "striking" as many complex patterns when seen againsta light background. In contrast to the "basic pattern", "ALL DARK" mayindeed be expressed flash-like. Similarly "ALL LIGHT" tends to be expressedvery briefly. Clearly, they do not appear in the most active phase of prey capture,but when expressed very briefly, "ALL DARK" and "ALL LIGHT" might havea startling effect similar to what has been suggested above for phase 2/3" (Mauris 1989) Mauris 1989 0 0 2 "indeed all loliginid are gregarious, from advanced juvenile stages onward, whereas Sepiola is certainly not gregarious at subadult and adult stages. However, personal field observations have provided evidence that very young Sepiola may form rather dense assemblies in the absence of any space limitation.Considering the rich pattern repertoire of early juveniles the question of intraspecific communication of post-hatching stages should at least be raised” (Mauris 1989) Mauris 1989 0 "indeed all loliginid are gregarious, from advanced juvenile stages onward, whereas Sepiola is certainly not gregarious at subadult and adult stages. However, personal field observations have provided evidence that very young Sepiola may form rather dense assemblies in the absence of any space limitation.Considering the rich pattern repertoire of early juveniles the question of intraspecific communication of post-hatching stages should at least be raised” (Mauris 1989) Mauris 1989 3 0 3 ? “the repertoire of colour patterns and body postures observed in Sepiola is relatively rich and appears particularly varied considering the absence of two important elements of patterning known in Sepia and Octopus…personal field observations have provided evidence that very young Sepiola may form rather dense assemblies in the absence of any space limitation. Considering the rich pattern repertoire of early juveniles the question of intraspecific communication of post-hatching stages should at least be raised” (Mauris 1989) Mauris 1989 1 "In particular, Rodrigues et. al. (2009) state that, indeed, it is the female that initiates mating by hovering over the male; a stance that is reported by all authors. In our opinion, the female while hovering might possibly send light cues to the male indicating her presence and availability to mate; in fact, according to the present results, the male fixed his eyes on the female for a few seconds before approaching her…Mating in Sepiolinae appears to occur preferentially during the hours of darkness, which is a further support to the hypothesis that the female visceral luminous glands might play a role in triggering the male response in species with luminous organs." (Bello & Deickert 2021) (Bello & Deickert 2021) Unsure evidence “Mauris (1988), in her thesis, provided detailed observations on some mating events in captive Sepiola affinis, all of them occurring at night. She wrote that the male grabs the female’s neck by his third arms, which are short and thicker than the other arms; the male’s third arms are also thicker than the female’s third arms; both his frst arms are inserted into the female mantle cavity; his second arms, which show to be very fexible, are lengthened and placed on the ventral side of the female’s mantle; his fourth arms are placed under the female’s head. The female’s mantle is at an angle of about 45° with respect to the male plane and in this position the pair at frst swims around and after a while, keeping on mating, go to rest on the bottom; here the female leans her arms on the ground. Mauris (1988) also discussed the behaviour of a male and a female of the same species placed together in a tank, where the female went to the bottom in a corner and kept still with her chromatophores expanded. Meanwhile, the male restlessly swam around her, keeping at about 5–6 cm distance, alternating swimming with staying still on the bottom of the tank with his arms spread out; this author herself was not able to judge, by just one observation, whether this behaviour might be deemed a courtship.” (Bello & Deickert 2021) 14 16 66.906 30.996 30 19 W-W
Sepiola robusta Sepiola robusta 178 "Depth range from 20 to 178 m" (Jereb & Roper, 2005) Jereb & Roper, 2005 15 "has been found at depths slightly over 150 m, but it is typically abundant in shallow waters (15–30 m)," (Jereb & Roper, 2005) Jereb & Roper, 2005 2 "Outer shelf." (Jereb & Roper, 2005) Jereb & Roper, 2005 15 45 45 N to 30 N (Jereb & Roper, 2005) (Jereb & Roper, 2005) 30 45 N to 30 N (Jereb & Roper, 2005) (Jereb & Roper, 2005) (Salman et al. 1998) 1 273.9375 "The normal life cycle of Sepiola robusta lasts between 6 and 9 months" (Boyle, 1983). (Boyle, 1983) 182.625 "The normal life cycle of Sepiola robusta lasts between 6 and 9 months" (Boyle, 1983). (Boyle, 1983) 1 1 1 in lab, sepiolines generally, “After leaving the sand, the animals approach their prey, arms placed together and stretched out forming an elongated cone which points towards the prey (they also circle round the prey in order to attack it from the side or from behind). Sepiolinae then dart swiftly (propelled by the funnel jet) on the prey and seize it, with their tentacles shooting forward simultaneously. The captured animal is pulled back, to be grabbed instantly by the arms. Since the prey is often larger than the attacking Sepiola or Sepielta, the predator may be dragged about wildly by its struggling prey for several seconds before the latter dies. Sepiolinae usually keep swimming when eating a very large prey, bat settle on the bottom (without burrowing) to devour prey of smaller size” (von Boletzky et al. 1971) von Boletzky et al. 1971 3 2 1 lab “Sand burrowing is common to all the species reared. Newly hatched animals, as well as those kept in a tank without sand for weeks, burrow when offered sand…Hiding in the sand is evidently a means of protection against predators; it does not serve for outwitting prey, since sepiolids do not capture prey passing by while buried in the sand. In all cases, they first leave the sand, and then attack the prey; they do not burrow again before they have finished feeding (provided they are not disturbed).” (von Boletzky et al. 1971) von Boletzky et al. 1971 1 "The fact that both S. atlantica and S. robusta, as well as other Sepiola species and Rondeletiola minor, may leave the bottom and move towards a light source, certaintly suggests the counter-illuminating function of their luminous glands" (Bello & Biagi 1995) Bello & Biagi 1995 1 Present (Jereb & Roper, 2005) (Jereb & Roper, 2005) 3 "As far as laboratory observations go. Sepiola robusm readily feeds on mysid shrimp at early juvenile stages, and on mysids, palaemonids and crangonids at advanced juvenile and adult stages (Fig. 4.6)" (Boyle, 1983). Boyle 1983 0 0 1 0 not gregarious (inferred from photo and video material) 1 0 1 1 "Males have been observed to guard females during courtship." (Jereb & Roper, 2005) (Jereb & Roper, 2005) (Hanlon & Messenger, 2018, p. 153) 17 16 47.104 32.016 24 20 W-W
Sepiola rondeletii Sepiola rondeletii 498 "As for Sepiola robusta, the bathymetric distribution range observed for males in the Strait of Sicily
(maximum depth recorded being 498 m) is definitely wider than that generally reported for the species
(Belcari et al., 1989; Orsi Relini and Bertuletti, 1989;
Guerra, 1992; Sartor and Belcari, 1995; Volpi et al.,
1995). However, considering another rather atypical
record recently mentioned for the lower Tyrrhenian
Sea (Wurtz et al., 1995), it seems likely that the
depth distribution of this species should be extended" (Jereb et al. 1997)		 Jereb & Roper, 2005 26 "Depth range from 26 to 498 m." (Jereb & Roper, 2005) Jereb & Roper, 2005 2 "Sandy and muddy substrates, common in Posidonia seagrass beds down to 35 m. Epibenthic, or mesobenthic," (Jereb & Roper, 2005) Jereb & Roper, 2005
Bello, 2019		 51 63 63 N to 12 N (Jereb & Roper, 2005) (Jereb & Roper, 2005) 12 63 N to 12 N (Jereb & Roper, 2005) (Jereb & Roper, 2005) 1 547.9 "longevity estimated at 18 months." (Jereb & Roper, 2005) (Jereb & Roper, 2005) 547.9 "longevity estimated at 18 months." (Jereb & Roper, 2005) (Jereb & Roper, 2005) NA 0 2 1 1 lab “Sand burrowing is common to all the species reared. Newly hatched animals, as well as those kept in a tank without sand for weeks, burrow when offered sand…Hiding in the sand is evidently a means of protection against predators; it does not serve for outwitting prey, since sepiolids do not capture prey passing by while buried in the sand. In all cases, they first leave the sand, and then attack the prey; they do not burrow again before they have finished feeding (provided they are not disturbed).” (von Boletzky et al. 1971) von Boletzky et al. 1971) 1 (Jereb & Roper, 2005) 2 "The species feeds on crustacea and small fishes." (Jereb & Roper, 2005) (Jereb & Roper, 2005) 0 0 2 0 not gregarious (inferred from photo and video material) NA 0 NA 34 13 58.89 36.27 18.2 27 21 23 W-W-NY
Sepioteuthis lessoniana Sepioteuthis lessoniana 100 "occurs from the surface to about 100 m depth throughout its area of distribution. " (Jereb & Roper, 2010) Jereb & Roper, 2010
Arkhipkin et al., 2015
Iglesias et al., 2014
Elsevier, 2014
Pratoomchat et al., 2011		 0 "occurs from the surface to about 100 m depth throughout its area of distribution. " (Jereb & Roper, 2010) Jereb & Roper, 2010
Arkhipkin et al., 2015
Iglesias et al., 2014
Elsevier, 2014
Pratoomchat et al., 2011		 2 **edge case** "In nature, S. lessoniana occurs inshore, near temperate rock reefs or coral reefs, and this may partially explain its adaptability to tanks (Segawa, 1987; Jackson, 1990)" (Lee et al., 1994). "Sepioteuthis lessoniana is a demersal neritic species that inhabits coral and rock reefs, seaweed, sea grass beds, and estuaries" (Nabhitabhata & Ikeda 2014) Lee et al., 1994; Nabhitabhata & Ikeda 2014) 99 52 52 N to 47 S (Jereb & Roper, 2010) Jereb & Roper, 2010) -47 52 N to 47 S (Jereb & Roper, 2010) Jereb & Roper, 2010) 1 1 "it consistently migrates inshore after winter to start mating and spawning in spring." (Jereb & Roper, 2010) (Jereb & Roper, 2010) (Arkhipkin et al., 2015) 1 “The bigfin squid preferred medium level of brightness during the first 6 hrs and then gradually changed to prefer low level of brightness at night…interpreted as innate migration to greater depth. Hence, the brightness of substrata might be associated with visual discrimination of depth by the squid. Moreover, the degree of preference that lowered to nonsignificant difference during the nighttime and might relate to photopositive taxis behaviour in low light environment, i.e. diurnal migration (Nabhitabhata 1978b)…congregate in shallow water during the daytime and migrate offshore to deeper water at night” (Nabhitabhata & Nilaphat 2000) Nabhitabhata & Nilaphat 2000
Segawa, 1995		 365.25 "Early length frequency analysis and field observations indicated a lifespan between 1 and 3 years for S. lessoniaNA(Jereb and Roper, 2006). However, rearing experiments and direct ageing techniques indicate a much shorter lifespan of » 6 months (Jackson and Moltschaniwskyj, 2002), with animals reaching sexual maturity between 110 and 140 days (Jereb and Roper, 2006). Significant variation in growth rates and maturity exist between equatorial, tropical, and
subtropical Indo-Pacific populations (Jackson and Moltschaniwskyj, 2002)." (Arkhipkin et al., 2015)		 182.625 "Early length frequency analysis and field observations indicated a lifespan between 1 and 3 years for S. lessoniaNA(Jereb and Roper, 2006). However, rearing experiments and direct ageing techniques indicate a much shorter lifespan of » 6 months (Jackson and Moltschaniwskyj, 2002), with animals reaching sexual maturity between 110 and 140 days (Jereb and Roper, 2006). Significant variation in growth rates and maturity exist between equatorial, tropical, and
subtropical Indo-Pacific populations (Jackson and Moltschaniwskyj, 2002)." (Arkhipkin et al., 2015)		 140 "Early length frequency analysis and field observations indicated a lifespan between 1 and 3 years for S. lessoniaNA(Jereb and Roper, 2006). However, rearing experiments and direct ageing techniques indicate a much shorter lifespan of » 6 months (Jackson and Moltschaniwskyj, 2002), with animals reaching sexual maturity between 110 and 140 days (Jereb and Roper, 2006). Significant variation in growth rates and maturity exist between equatorial, tropical, and subtropical Indo-Pacific populations (Jackson and Moltschaniwskyj, 2002)." (Arkhipkin et al., 2015) (Arkhipkin et al., 2015) 110 "Early length frequency analysis and field observations indicated a lifespan between 1 and 3 years for S. lessoniaNA(Jereb and Roper, 2006). However, rearing experiments and direct ageing techniques indicate a much shorter lifespan of » 6 months (Jackson and Moltschaniwskyj, 2002), with animals reaching sexual maturity between 110 and 140 days (Jereb and Roper, 2006). Significant variation in growth rates and maturity exist between equatorial, tropical, and
subtropical Indo-Pacific populations (Jackson and Moltschaniwskyj, 2002)." (Arkhipkin et al., 2015)		 (Arkhipkin et al., 2015) 2 1 1 S. lessoniaNAbegan to exhibit the tentacular strike attack after 30 days of age" (Sugimoto & Ikeda, 2013). Sugimoto & Ikeda, 2013 1 “Both species showed alterations in their choice of body pattern during the attack sequence at elevated CO 2; while there was a clear preference for the uniform blanch body display in controls, there was a significant increase in the dark mottle pattern in squid exposed to high CO 2…When squid attacked with the uniform blanch body pattern, their transparent appearance much more easily blended in with the background shade.” (Spady et al. 2018) Spady et al. 2018 2 2 1 “In addition to its appearance, for a squid, the size variation among schoolmates and the number of schoolmates relates directly to the functions of a school… From the principle of the oddity effect of groups (i.e. predators may preferentially target the least common phenotype in a group), evolutionary pressure for grouping with similar individuals is produced (Krause & Ruxton, 2002)…Smaller S. lessoniaNA(ML less than 100 mm) form a small school (schools of 10 and 20 squid) and exhibit narrow size-variation among schoolmates. In contrast, larger S. lessoniaNA(ML more than 100 mm) tend to form a large school (schools of 40 and 100 squid) with various-sized schoolmates. In a school with 100 squid, the size-range of the schoolmates is wider than that in a school with 40 squid” (Sugimoto et al. 2013) Sugimoto et al. 2013 1 "Among teuthoids, only the loliginids Sepioteuthis sepioidea and S. lessoniaNAshow a ‘typical’ Deimatic Display, with false eyespots on the mantle and a pale or lightly mottled coloration. Lolliguncula brevis (Dubas et al., Reference Dubas, Hanlon, Ferguson and Pinsker1986) and L. panamensis (Moynihan & Rodaniche, Reference Moynihan and Rodaniche1982) tend to show displays intermediate between the Deimatic and the Flamboyant (described below); that is, they have arms flared in Upward V-curl, light bodies with some sort of dark border (on the fins or the flared arms), but they do not have conspicuous false eyespots." (Hanlon & Messenger, 2018, p. 125) Hanlon & Messenger, 2018, p. 125 7 "In the natural environment this species preys primarily on prawns and fishes, occasionally on stomatopods and crabs." (Jereb & Roper, 2010) Jereb and Roper 2010
Igelsias et al 2014
Moltschaniwskyj & Jackson, 2000		 0 0 3 2 “(stomach) long-beaked common dolphin (Delphinus capensis)” (Ahn et al., 2014) Ahn et al., 2014
Amir et al., 2005		 1 3 schooling/shoaling behavior: Okinawa Island "16 schools of S. lessoniaNAwere observed, containing 8 to more than 100 squid of sizes ranging from 50 to 200 mm ML (Table 2). Squid schools were divided into 4 categories as containing 100 squid, 40 squid, 20 squid and 10 squid. SCHOOLS 1 through to 4 contained different sizes of squid (100 to 200 mm ML for SCHOOL 1; 150 to 200 mm ML for SCHOOLS 2, 3 and 4). SCHOOLS 5 through to 16 mainly contained squid of the same size (50, 100, or 150 mm ML)" (Sugimoto et al. 2016) Sugimoto et al. 2013, 2016 1 schooling/shoaling behavior: Okinawa Island "16 schools of S. lessoniaNAwere observed, containing 8 to more than 100 squid of sizes ranging from 50 to 200 mm ML (Table 2). Squid schools were divided into 4 categories as containing 100 squid, 40 squid, 20 squid and 10 squid. SCHOOLS 1 through to 4 contained different sizes of squid (100 to 200 mm ML for SCHOOL 1; 150 to 200 mm ML for SCHOOLS 2, 3 and 4). SCHOOLS 5 through to 16 mainly contained squid of the same size (50, 100, or 150 mm ML)" (Sugimoto et al. 2016) Sugimoto et al. 2013, 2016 9 0 9 1 “Although the shape is similar, the belt-shaped appearance of S. lessoniaNAschools also has a unique characteristic. Unlike birds and dolphins, S. lessoniaNAcan hover in mid-water. Hence, hydrodynamic advantages are an unlikely reason for this and other static appearances, despite the difference in the buoyancy and density effect between air and water. Instead, visual contact might be used for communication between school members, since squid possess a highly developed and laterally positioned lens and can change their body patterns. These physical attributes may suggest that squid have visual communication with members of the school.” (Sugimoto et al. 2013) Sugimoto et al. 2013 1 squids raised in captivity: “animals sometimes accentuated the visibility of their gonads (‘Accentuated Gonads’) so that their gonads appeared bright white through their mantles, a behaviour described as a “chromatic signal” by HANLON & MESSENGER (1996, p. 125). Accentuated Gonads was usually, although not always, reciprocated by an adjacent animal…. The more sexually active individuals (more often involved in what appeared to be reproductive behaviour), both male and female, were more likely to show Accentuated Gonads than were the less sexually active individuals. Accentuated Gonads was observed in situations that were not clearly social, suggesting that this signal could simply indicate the sender’s reproductive condtion. In support for this hypothesis, the frequency of Accentuated Gonads was positively correlated with the frequency of receiving a reproductive event.” (Boal & Gonzalez 1998) Boal & Gonzalez 1998 6 Total number of body patterns is 6 (Hanlon & Messenger, 2018) Hanlon & Messenger, 2018 1 "In the waters of India, these squid migrate inshore after winter to begin mating and spawning (Jereb and Roper, 2006)" (Arkhipkin et al., 2015) Arkhipkin et al., 2015 1 Squids raised in lab: “Males directed ‘reproductive’ behaviour towards each other as well as towards females. In 31 of the 63 observed events in which the recipient was identified with confidence, the recipient was female, while in 32 cases the recipient was male…The likelihood that a reproductive event would reach Contact was related to the sex of the recipient. Of the 32 approaches to male recipients, 13 ended with Contact; for female recipients, 3 of the 31 approaches ended with Contact…Interactions with females were more likely to be terminated before apparent spermatophore transfer…We hypothesize that males do have a bias towards female recipients but were disproportionately rebuffed by them…among squids cultured in the laboratory, males usually mature sexually before females (personal observation). Medium and large males initiated most reproductive events, indicating that they were sexually mature. Females, on the other hand, did not spawn for at least another month after this particular subset of data was collected so females may have been sexually immature” (Boal & Gonzalez 1998) Boal & Gonzalez 1998
Lin and Chiao 2017
Lin et al 2017		 1 raised in captivity: “squids sometimes darkened their mantle and spread their arms widely (‘Spread Arms’), at times in a slightly head-down and arms-up, V-position. Spread Arms was often shown while following or chasing another animal. This behaviour seems not unlike the Zebra Display of Sepzotezlthis sepiozdea (MOYNIHAN et al. 1982; MOYNIHAN 1985; HANLON & MESSENGER 1996) but was less dramatic in body coloration…Spread Arms was also used by males when near a mate or potential mate ” (Boal & Gonzalez 1998) Boal & Gonzalez 1998 1 During mating, squid form close pairs, and paired males exhibit characteristic agonistic colour patterns against other intruding males" (Jereb & Roper, 2010) Jereb & Roper, 2010
Lin and Chiao 2017		 1 "The male used three different mating behaviors: male-parallel (MP), male-upturned (MU) and sneaking. Male competition over females frequently occurred before and during the female egg-laying period, and the outcome of most fights depended on male body size. Larger males guarded their partners from other males and performed MP mating during the egg-laying period of the paired females. In contrast, there was no pairing and mate guarding in MU mating and sneaking, which were adopted by smaller subordinate males as alternative tactics outside female egg-laying period and during the period, respectively. MP matings were 95% successful, but more than half of MU matings were unsuccessful. Higher mating success in MP mating was achieved through pairing, whereas males in MU mating were less successful because mating attempts without pair formation were often foiled by escape of the female. Sneaking was successful in all cases but occurred less frequently. Spermatophores were attached at the opening of the oviduct in MP mating, whereas they were attached around the female buccal membrane in MU mating and sneaking. Considering the route of egg transportation, higher fertilization success can be expected in MP mating because of the advantageous location of the attached spermatophores. Our results suggest that MP mating is used by larger, paired males during the female egg-laying period, and that MU mating and sneaking are alternative tactics adopted by smaller, subordinate males. These alternative mating behaviors would be conditional strategy dependent on relative body size, because some individual males displayed both MP and MU mating behaviors" (Wada et al., 2005). Wada et al., 2005
Hanlon & Messenger, 2018, p. 166		 1 "The male used three different mating behaviors: male-parallel (MP), male-upturned (MU) and sneaking. Male competition over females frequently occurred before and during the female egg-laying period, and the outcome of most fights depended on male body size. Larger males guarded their partners from other males and performed MP mating during the egg-laying period of the paired females. In contrast, there was no pairing and mate guarding in MU mating and sneaking, which were adopted by smaller subordinate males as alternative tactics outside female egg-laying period and during the period, respectively. MP matings were 95% successful, but more than half of MU matings were unsuccessful. Higher mating success in MP mating was achieved through pairing, whereas males in MU mating were less successful because mating attempts without pair formation were often foiled by escape of the female. Sneaking was successful in all cases but occurred less frequently. Spermatophores were attached at the opening of the oviduct in MP mating, whereas they were attached around the female buccal membrane in MU mating and sneaking. Considering the route of egg transportation, higher fertilization success can be expected in MP mating because of the advantageous location of the attached spermatophores. Our results suggest that MP mating is used by larger, paired males during the female egg-laying period, and that MU mating and sneaking are alternative tactics adopted by smaller, subordinate males. These alternative mating behaviors would be conditional strategy dependent on relative body size, because some individual males displayed both MP and MU mating behaviors" (Wada et al., 2005). Wada et al., 2005
Jones et al 2019
Apostolico and Marian 2019		 1 "Mating pairs touch their newly laid egg capsules and stay near them to guard them from other pairs (Fig. 17.12). On average, mating pairs touch their eggs for approximately 6 s and guard them for 45 s. Similar behaviour is observed in both cultured (Lee et al. 1994) and wild squid (Segawa 1987)" (Iglesias et al., 2014). Iglesias et al., 2014 220 112 582.38 113 C20
Sepioteuthis sepioidea Sepioteuthis sepioidea 20 "most common in waters of 3-7 m depth, although divers have reported it from 20 m (LaRoe, 1967)." (Cairns 1976) (Cairns, 1976; Jereb & Roper, 2010; Haimovici et al., 1989; Boycott 1965; Anchietta et al. 2007) 0 "It occurs at depths of 0 to 20 m, mostly 3
to 7 m." (Jereb & Roper, 2010)		 (Cairns, 1976; Jereb & Roper, 2010; Haimovici et al., 1989; Boycott 1965; Anchietta et al. 2007) 2 **edge case** JM: should maybe be pelagic. "S. sepioidea is a shallow water, inshore species most common in waters of 3-7 m depth, although divers have reported it from 20 m (LaRoe, 1967)" (Cairns, 1976) (Cairns, 1976) 41 28 28 N to 13 S (Jereb & Roper, 2010) even though "A single record from the waters off Woods Hole, Massachussets (41°32’N, 70°41’W; Mercer, 1970b), conspicuously far from the normal
northern distribution limit of the species, was explained as an example of transportation by the Gulf Stream. A single record from
southeastern Brazil waters off Buzios (23°47’S; 45°10’W) also exists (Begossi and Duarte, 1988); the specimen was associated
with a rocky and sandy substrate, at about 5 to 6 m deep, in March 1987. However, the species was not reported in the Brazilian
waters afterwards, till the recent observations off northeastern Brazil (Nunes de Anchieta et al., 2007)." (Jereb & Roper, 2010)		 (Jereb & Roper, 2010) -13 28 N to 13 S (Jereb & Roper, 2010) even though "A single record from the waters off Woods Hole, Massachussets (41°32’N, 70°41’W; Mercer, 1970b), conspicuously far from the normal
northern distribution limit of the species, was explained as an example of transportation by the Gulf Stream. A single record from
southeastern Brazil waters off Buzios (23°47’S; 45°10’W) also exists (Begossi and Duarte, 1988); the specimen was associated
with a rocky and sandy substrate, at about 5 to 6 m deep, in March 1987. However, the species was not reported in the Brazilian
waters afterwards, till the recent observations off northeastern Brazil (Nunes de Anchieta et al., 2007)." (Jereb & Roper, 2010)		 (Jereb & Roper, 2010) 1 487 Life span: 12-16 months (Mather, 2016) (LaRoe 1971; Mather, 2016) 365.3 ~1 year lifespan (LaRoe 1971) (LaRoe 1971; Mather, 2016) 182.6 based on specimens raised in lab: "From the growth and maturity data here, it appears that Sepioteuthis sepioidea, at least, is mature and can reproduce at an age of 6 months" (LaRoe, 1971) (LaRoe, 1971) 182.6 based on specimens raised in lab: "From the growth and maturity data here, it appears that Sepioteuthis sepioidea, at least, is mature and can reproduce at an age of 6 months" (LaRoe, 1971) (LaRoe, 1971) 6 5 1 "Moynihan and Rodaniche (1982) described how Sepioteuthis sepioidea mimics sargassum weed as it hunts. This involves the individual making the appropriate locomotor, postural, textural and chromatic adjustments, and also associating itself with clumps of weed, from which it can dart out at prey and to which it will subsequently return (Fig. 4.llb; see also §4.4.l)." (Hanlon & Messenger, 2018, p. 85) Hanlon & Messenger, 2018, p. 85
Hanlon & Messenger, 2018, p. 89		 1 "The loliginid squid Sepioteuthis sepioidea wave their dorsal arm pairs from side to side before attacking prey, which may indicate hypnotizing or luring." (Hoving et al 2013) Hoving et al 2013
Hanlon & Messenger, 2018, p. 83		 0 "It is especially interesting that there is no sign of cooperative hunting by the highly developed Caribbean reef squid, Sepiotertthis sepioidea. As Moynihan and Rodaniche (1982, p. 23] make clear, ‘the behavior of one individual may alert its companions to the presence of possible prey in the neighborhood. The members of a group do not, however, seem to cooperate to drive, chase, or trap their prey by any purposeful communal effort.’" (Hanlon & Messenger, 2018, p. 86) Hanlon & Messenger, 2018, p. 86 1 "The squid Sepioteurhis sepioidea (and many decapodiformes) also stalk fish and shrimp (Fig. 4.11a), mostly at night [Forsythe & Hanlon. unpublished data);" (Hanlon & Messenger, 2018, p. 83) Hanlon & Messenger, 2018, p. 83
Hanlon & Messenger, 2018, p. 89		 1 The squid Sepioteuthis sepioidea may also engage in a form of speculative hunting" (Hanlon & Messenger, 2018, p. 84) Hanlon & Messenger, 2018, p. 84 1 "A squid makes an anterior-first Jerk or jets to the appropriate distance from a prey item, in down Tilt if the prey is on the substrate. Then it Maintains with fin Undulate and low-velocity Jets, sometimes using one or more components of Sway. Next the arms quickly Splay and tentacles Strike. They Contract and the arms can Grasp and the squid feed (note the approaching and selection before this stereotyped prey capture may be variable, see Hanlon and Messenger 1996)." (Mather et al. 2010) Mather et al. 2010
Hanlon & Messenger, 2018, p. 82		 1 "Hanlon and Forsythe (unpublished) have filmed Sepioteuthis sepioidea swimming backwards while displaying two false eyespots, so that it looked very like a herbivorous parrotfish common in the habitat, especially as the arms and tentacles were held together and waved from side-to-side like the caudal fin of a fish. Swimming in this way, the squid swam amidst small reef ish and occasionally made short forward thrusts to grasp fish (Fig. 4.l]d]." (Hanlon & Messenger, 2018, p. 85) Hanlon & Messenger, 2018, p. 85 11 10 1 "Since these squid do not rest on the bottom, their primary defence cannot be hiding on or in the substrate like octopuses and cuttlefish. During the day, this species shoals over seagrass beds adjacent to the reefs, where there are fewer predators. They maintain countershading and translucency when in the water column, and this form of crypsis can be very effective (Fig. 5.3c), yet they have many body patterns to help conceal themselves amidst seagrasses, corals and sponges (Fig. 5.3d; Fig. 5.13g,h). For much of the time the squid are in linear shoals showing a dark body pattern (termed Basic by Moynihan & Rodaniche, Reference Moynihan and Rodaniche1982) that is not cryptic. They have apparent ‘sentinels’ in each shoal, and usually there are squid facing in all directions at any one time (Fig. 5.24). Thus, primary defences are (1) shoaling with some squid facing in all directions but not in apparently cryptic coloration, or (2) when single or in pairs, deploying cryptic body patterns for background matching, disruptive patterning and masquerade." (Hanlon & Messenger, 2018, p. 138) (Hanlon & Messenger, 2018, p. 138) 1 "Since these squid do not rest on the bottom, their primary defence cannot be hiding on or in the substrate like octopuses and cuttlefish. During the day, this species shoals over seagrass beds adjacent to the reefs, where there are fewer predators. They maintain countershading and translucency when in the water column, and this form of crypsis can be very effective (Fig. 5.3c), yet they have many body patterns to help conceal themselves amidst seagrasses, corals and sponges (Fig. 5.3d; Fig. 5.13g,h). For much of the time the squid are in linear shoals showing a dark body pattern (termed Basic by Moynihan & Rodaniche, Reference Moynihan and Rodaniche1982) that is not cryptic. They have apparent ‘sentinels’ in each shoal, and usually there are squid facing in all directions at any one time (Fig. 5.24). Thus, primary defences are (1) shoaling with some squid facing in all directions but not in apparently cryptic coloration, or (2) when single or in pairs, deploying cryptic body patterns for background matching, disruptive patterning and masquerade." (Hanlon & Messenger, 2018, p. 138) (Hanlon & Messenger, 2018, p. 138) 1 "Since these squid do not rest on the bottom, their primary defence cannot be hiding on or in the substrate like octopuses and cuttlefish. During the day, this species shoals over seagrass beds adjacent to the reefs, where there are fewer predators. They maintain countershading and translucency when in the water column, and this form of crypsis can be very effective (Fig. 5.3c), yet they have many body patterns to help conceal themselves amidst seagrasses, corals and sponges (Fig. 5.3d; Fig. 5.13g,h). For much of the time the squid are in linear shoals showing a dark body pattern (termed Basic by Moynihan & Rodaniche, Reference Moynihan and Rodaniche1982) that is not cryptic. They have apparent ‘sentinels’ in each shoal, and usually there are squid facing in all directions at any one time (Fig. 5.24). Thus, primary defences are (1) shoaling with some squid facing in all directions but not in apparently cryptic coloration, or (2) when single or in pairs, deploying cryptic body patterns for background matching, disruptive patterning and masquerade." (Hanlon & Messenger, 2018, p. 138) (Hanlon & Messenger, 2018, p. 138) 1 masquerade (Hanlon & Messenger, 2018, p. 114) (Hanlon & Messenger, 2018, p. 114) 1 Mimicry: "Caribbean reef squid have also been observed using defensive mimicry, exhibiting body patterns that strikingly resemble parrot fish, Scarus taenipterus, found on the same reef (Moynihan & Rodaniche, 1982)." (Schnell et al., 2020) (Schnell et al., 2020) 1 "Since these squid do not rest on the bottom, their primary defence cannot be hiding on or in the substrate like octopuses and cuttlefish. During the day, this species shoals over seagrass beds adjacent to the reefs, where there are fewer predators. They maintain countershading and translucency when in the water column, and this form of crypsis can be very effective (Fig. 5.3c), yet they have many body patterns to help conceal themselves amidst seagrasses, corals and sponges (Fig. 5.3d; Fig. 5.13g,h). For much of the time the squid are in linear shoals showing a dark body pattern (termed Basic by Moynihan & Rodaniche, Reference Moynihan and Rodaniche1982) that is not cryptic. They have apparent ‘sentinels’ in each shoal, and usually there are squid facing in all directions at any one time (Fig. 5.24). Thus, primary defences are (1) shoaling with some squid facing in all directions but not in apparently cryptic coloration, or (2) when single or in pairs, deploying cryptic body patterns for background matching, disruptive patterning and masquerade." (Hanlon & Messenger, 2018, p. 138) (Hanlon & Messenger, 2018, p. 138) 1 "Among teuthoids, only the loliginids Sepioteuthis sepioidea and S. lessoniaNAshow a ‘typical’ Deimatic Display, with false eyespots on the mantle and a pale or lightly mottled coloration. Lolliguncula brevis (Dubas et al., Reference Dubas, Hanlon, Ferguson and Pinsker1986) and L. panamensis (Moynihan & Rodaniche, Reference Moynihan and Rodaniche1982) tend to show displays intermediate between the Deimatic and the Flamboyant (described below); that is, they have arms flared in Upward V-curl, light bodies with some sort of dark border (on the fins or the flared arms), but they do not have conspicuous false eyespots." (Hanlon & Messenger, 2018, p. 125) (Hanlon & Messenger, 2018, p. 125) (Boycott 1965) 1 “Our experimental study tested the hypothesis that squid use ink as a
defense against attacks by an ecologically relevant fish predator. We
examined the effects of ink from the Caribbean reef squid, S. sepioidea,
on the behavior of juvenile French grunts, H. flavolineatum, in three
sets of experiments. In one set, an ink pseudomorph released between the
food and fish significantly changed the fish's behavioral responses,
causing avoidance of or biting at the ink pseudomorph and a delay in
time to capture the food. In the second set of experiments, ink added to
a meat-flavored paper disc reduced its palatability as measured by
handling and acceptance. In the third set, ink added to an otherwise
unflavored disc did not increase the disc's palatability as measured by
handling and acceptance… ours is the first experimental demonstration
that squid ink is an effective defense against predators” (Wood et al.
2010)		 (Wood et al.
2010)		 1 (Wood et al. 2010) 1 "The coral reef squid Sepioteuthis sepioidea uses rapid jetting as the only defence when its most dangerous predators approach" (Hanlon & Messenger, 2018, p. 124) (Hanlon & Messenger, 2018, p. 124) (Boycott 1965) 1 "‘Flying squid’ (Fig. 5.20) are well known and particularly common among oceanic oegopsids that live near the surface. One of the best documented accounts is that of Muramatsu et al. (Reference Muramatsu, Yamamoto and Abe2013) who filmed a school of ca. 100 ommastrephids ["very likely young of Ommastrephes bartramii or possibly S. oualaniensis on the basis of habitat and morphology"; Muramatsu, Yamamoto and Abe 2013:1173] simultaneously exit the water for about 3 seconds, then continue to jet water when airborne and to glide for more than 30 m (individual speeds of 9–11 m s–1). The gliding was accomplished by spreading the fins and the arms, the latter of which have a thin membrane that increases the ‘wing surface’ and provides more lift than the fins (Azuma, Reference Azuma2006). Macia et al. (Reference Macia, Robinson, Craze, Dalton and Thomas2004) provided new data on Sepioteuthis sepioidea flying and reviewed many reports of flying squid (e.g. Arata, Reference Arata1954; Cole & Gilbert, Reference 303Cole and Gilbert1970; Packard, Reference Packard1972; Azuma, Reference Azuma1981). Rather than simple gliding (as previously thought) after incidental exit from the water, this behaviour involved jet propulsion, generation of lift force (arms and fins), and control of different body postures in different phases of flight. This form of protean escape is likely to be effective against their aquatic visual predators. Although impressive, squid do not glide nearly as far as flying fish." (Hanlon & Messenger, 2018, p. 124) (Hanlon & Messenger, 2018, p. 124) 1 To less intense threats, squid showed an equally large variety of body patterns. Moynihan & Rodaniche (Reference Moynihan and Rodaniche1982) reported that they saw 279 permutations of seven components (not specified) for all types of behaviour over their lengthy field study; even allowing for misinterpretation, it is clear that this species can show many patterns. We estimate from our own extensive field observations and video recordings that Sepioteuthis can show perhaps several dozen body patterns to predators during defence interactions. These tactics are effective: no successful predation – and only a few unsuccessful attacks – has ever been seen during many hundreds of hours of observation by divers and hundreds of approaches by predators (Hanlon & Forsythe, unpublished data; Mather, Reference Mather2010). Thus, the many body patterns serve two purposes in this context: they interrupt the approach/attack sequence of fish predators, and, as suggested by Moynihan and Rodaniche (Reference Moynihan and Rodaniche1982), they possibly aid conspecifics in recognising alarm signals from sentinels. The body patterns may also tell the predator that it has been detected. By any measure, the behaviour of Sepioteuthis sepioidea is sophisticated and highly evolved." (Hanlon & Messenger, 2018, p. 138) (Hanlon & Messenger, 2018, p. 138) 10 squid collected from Clearwater Beach and Gibbets Bay in Bermuda and kept in lab: "The squid were offered a continuous supply of live locally collected silversides: Hogmouth Fry Anchoa choerostoma, Blue Fry Jenkinsia lamprotaenia, Rush Fry Allanetta harringtonensis, Pilchard Harengula humeralisi, and Anchovy Sardinella anchovia. We have observed S. sepioidea feeding on these fish in the wild and they are readily accepted as food in the lab (Replinger and Wood, 2007). Diets were enriched daily with gammarid amphipods. A sporadic diet enrichment of juvenile Sergeant Majors, breams, small crabs and shrimp were also offered."(Zeeh & Wood 2009) Zeeh and Wood 2009
Jereb and Roper 2010		 0 0 2 6 "The common Caribbean reef squid, Sepioteuthis sepioidea, has been observed being
attacked by several teleosts: yellow jacks (Caranx bartholomaei), bar jacks
(Caranx ruber), mutton snapper (Lutjanis analis), cero mackerel (Scomberomoras
regalis), great barracuda (SphyraeNAbarracuda) and houndfish (Tylosurus
crocodilus) (Moynihan & Rodaniche, Reference Moynihan and Rodaniche1982;
Hanlon & Forsythe, unpublished data; Mather, Reference Mather2010). The
fish may approach from high or low in the water column from anywhere in the
360° field of view, sometimes slicing through the middle of the linearly
arranged squid shoal to break it up into small groups. Barracuda, in
particular, often remain motionless near the squid for long periods of time so
that the squid must remain vigilant towards them while simultaneously observing
other passing predators." (Hanlon
& Messenger, 2018, p. 145)		 (Hanlon & Messenger, 2018) 1 3 “Squid were almost always in groups in the daytime (Moynihan & Rodaniche 1982), but dispersed to hunt at night. Group location was similar over weeks but exact group membership varied somewhat from day to day, which could be a fission–fusion organization heavily influenced by predation (Mather 2010)…Squid were nearly obligate members of their schools, with a membership of around 5–15 at maturity and between 2 and 50 as young individuals. Subadult squid often formed an approximate linear arrangement with no specific position for any individual.” (Mather 2016) Mather 2016 1 “Squid were almost always in groups in the daytime (Moynihan & Rodaniche 1982), but dispersed to hunt at night. Group location was similar over weeks but exact group membership varied somewhat from day to day, which could be a fission–fusion organization heavily influenced by predation (Mather 2010)…Squid were nearly obligate members of their schools, with a membership of around 5–15 at maturity and between 2 and 50 as young individuals. Subadult squid often formed an approximate linear arrangement with no specific position for any individual.” (Mather 2016) Mather 2016 8 1 1 "Simultaneous dual signalling is occurring during these male displays (Fig. 6.14), which can be quite dynamic. The female is receiving a signal of the standard calm courtship coloration of uniform brown in the male, whereas approaching males receive a highly aggressive all bright white signal. If the male shows the Lateral Silver Display to the female, she is repelled and the male loses his mate. Thus, when the female moves to his other side, he must quickly change sides with his patterning, which is done very smoothly as illustrated in video frame grabs in Fig. 6.14a. Only two other species have been seen to perform this feat: Sepia latimanus (Corner & Moore, Reference Corner and Moore1980) and Sepia plangon (Fig. 6.14c; Brown, Garwood & Williamson, Reference Brown, Garwood and Williamson2012)." Hanlon & Messenger, 2018, p. 165-171 Hanlon & Messenger, 2018, p. 165-171 7 1 “Three distinct types of behaviour were observed: (A) fish (P. virginicus or H. aurolineatum) swimming along with squid in the centre of the group; (B) fish (H. aurolineatum, P. virginicus, H. steindachneri or C. bartholomaei) swimming along with squid at the edges of the group; and (C) fish (H. aurolineatum) ‘cutting’ the squid group in a wandering or ‘zigzag’ swimming pattern, in and out of the group on a regular basis… In all records squid seemed unconcerned about the presence of fish. Although S. sepioidea is known as a predator of small fish, no aggressive behaviour was recorded.” "We suggest that this behaviour is mainly related to protection for juvenile fish against potential pelagic predators." (Anchieta et al. 2007); "Most squid gather with conspecifics, though Moynihan and Rodaniche (1982) noted S. sepioidea swimming with Doryteuthis (Loligo) plei. Groups sort by size; although S. sepioidea are attracted to conspecifics what-
ever the size, smaller animals are at risk for cannibalism from larger ones and so maintain several body lengths distance. It has been suggested that squid on the end of a line are sentinels, watching for predators and escaping first from them (Hanlon and Messenger 1996)" (Iglesias et al., 2014)		 Anchieta et al. 2007; Iglesias et al., 2014 0 "Despite the lack of communal or visually stimulated spawning, there is some evidence of synchronous spawning. On two occasions, several clutches of eggs appeared in the same general area at the same time…In both cases, no egg clusters had been seen for several months prior to or after the one brief period. Such occurrences suggest that a school or schools may select an area for spawning, and that several individuals within the school spawn during the same period, although not contiguously. On one occasion, a large cluster found under one rock appeared to be divided into two separate basal groups. The different stages of development between the two groups indicated that they were laid from 10 to 14 days apart. Possibly two females happened to discover the same "hiding" place or perhaps a single female revisited the site for a second spawning." (LaRoe 1971a) (LaRoe 1971a) 1 " Males defended females from other males, particularly with an agonistic Zebra display. Male–female pairs exchanged Saddle-Stripe displays, after which males might display an on–off Flicker. There was considerable female choice. Only if a female responded to this display with a parallel Rocking action would she pair and would the males deposit spermatophores at the base of her arms, and only 50% of the time did females move the spermatophores internally to where sperm might be released and stored in the oviducal gland for later fertilization of eggs." (Mather 2016) & “Females initiated sexual interactions with Saddle displays. In 2000, 299 of these displays were observed, most at the same time as male Stripe and in a female-over position…These exchanges did not appear to be primarily for pair-maintenance, as 50 were observed <30 s after one male ‘took over’ consortship from another, 43 after a male– male Zebra exchanged challenge and 175 during stable pairing…Thus, the exchange was more pair-initiating than pair-maintaining…Saddle was seldom observed – three times in 1999 and once in 2000 – by animals known to be male. In each case, a smaller male was responding to a sexual display by a larger one. This display may have functioned in that situation as a device to de-escalate a contest.” (Mather 2016) (Mather 2016) animals captured off Bimini Islands and kept in pens: "Sexual behavior usually appeared to be initiated by an aggressive male who selected a female and swam parallel to her, fluttering his fins rapidly, and displaying a pale reddish color pattern, particularly visible on his ventral surface. Typically, the females showed little initial response, either paying little noticeable attention or avoiding the male. There were no signs of open defense but occasionally the females would jump out of the water when pursued by the males. Later, when the females swam parallel to the males and displayed their own courtship patterns the males assumed a broad dark brown band mid-dorsal on the mantle which was longitudinally bisected by a stripe of light iridiocytes…The courtship pattern of the females involved a generalized contraction of the chromatophores except in a region on the dorsal mantle. Here a mid-dorsal dark brown stripe on the anterior half of the mantle was crossed by a dark brown yoke which went about half way around the mantle …Copulation was seldom observed but in the instances when it was, it occurred extremely rapidly, with the animals paralleling each other. The seminal receptacles of eight females were examined after these behavior patterns had been observed and all were filled with spermatophores." (Arnold 1965) 1 " Males defended females from other males, particularly with an agonistic Zebra display. Male–female pairs exchanged Saddle-Stripe displays, after which males might display an on–off Flicker. There was considerable female choice. Only if a female responded to this display with a parallel Rocking action would she pair and would the males deposit spermatophores at the base of her arms, and only 50% of the time did females move the spermatophores internally to where sperm might be released and stored in the oviducal gland for later fertilization of eggs." (Mather 2016) (Mather 2016) 1 “Males often swam in consortship, interposed between a female and another male. During consortship, males usually displayed some area of Zebra to other males…As noted in Mather (2004), escalation to the Zebra Spread resulted in a ritualized visual exchange, maintained for up to minutes and in over–under positions, though with little to no physical contact (see Moynihan & Rodaniche 1982, Figure 27). These high-intensity displays were more likely after entry of a novel male into the group, when two consort pairs were close together, and especially if a pair within a group displayed Saddle-Stripe or one male displayed Flicker” (Mather 2016) (Mather 2016) 1 "Males defended females from other males, particularly with an agonistic Zebra display. Male–female pairs exchanged Saddle-Stripe displays, after which males might display an on–off Flicker. There was considerable female choice. Only if a female responded to this display with a parallel Rocking action would she pair and would the males deposit spermatophores at the base of her arms, and only 50% of the time did females move the spermatophores internally to where sperm might be released and stored in the oviducal gland for later fertilization of eggs." (Mather 2016) (Mather 2016) 1 Venezuela coast "While two consorts were exchanging Zebra displays, a subordinate Sneaker male might give a Flicker to an unguarded female, and mating could follow…Subordinate males stayed and acted as a sneaker until they matured, or left the group" (Mather 2016) (Mather 2016) "the loliginid squid S. sepioidea, in which early adult males play sneaker tactics, but later change to consort tactics as they grow (Mather, 2016)." (Apostolico and Marian 2019) 0 "Females lay eggs in protected areas on the sea bottom but give no parental care" (Mather 2010) (Mather 2010) 0 "Females lay eggs in protected areas on the sea bottom but give no parental care" (Mather 2010) (Mather 2010) 74 58 261.2 150 NY
Spirula spirula Spirula spirula 1000 [AL: A lot of older records say that it presumably lives deeper, but later sources seem not to find any adults deeper than 800-1000] "Clarke has shown that in the day Spirula is
centred around 600-700 m, with occasional animals going down to depths of about
1000 m. Our measurements show that the shells of Spirula in the intact animal can stand
pressure found at depths around 1700 m. The lowest pressure at which a shell imploded
corresponded to a depth of about 1300 m" (Denton & Gilpin-Brown, 1971)		 (Lukeneder et al., 2008; Cairns, 1976; Cairns, 1976; Jereb & Roper, 2005; Voss, 1956; Mutvei, 2017; Lukeneder, 2016; Ohkouchi et al., 2013; Price et al., 2009; Lukeneder et al., 2008; Denton & Gilpin-Brown, 1971; Clarke, 1986; Lindsay et al. 2020; Roper & Young, 1975) 100 "It has not been observed or filmed in its
habitat, but is presumed to inhabit mesopelagic
(550–1,000 m deep) waters of tropical and subtropical
Atlantic and Indo-West Pacific regions (Norman 2,000).
The species is thought to live in groups and exhibits diel
vertical migration, based on evidence using sampling with
a closing net (Clarke 1969). During the day, it stays at
550–1,000 m depth and then rises to feed at 100–300 m
depth during the night. Spawning has been suggested to
occur in deep water close to the seafloor (Bruun 1943;
Nesis 1987), a hypothesis supported by the observation of
juveniles (0.5 cm in length) at depths of 1,000–1,750 m
(Clarke 1970)." (Ohkouchi et al., 2013)		 (Lukeneder et al., 2008; Cairns, 1976; Cairns, 1976; Jereb & Roper, 2005; Voss, 1956; Mutvei, 2017; Lukeneder, 2016; Ohkouchi et al., 2013; Price et al., 2009; Lukeneder et al., 2008; Denton & Gilpin-Brown, 1971; Clarke, 1986; Lindsay et al. 2020; Roper & Young, 1975) 1 "It has not been observed or filmed in its
habitat, but is presumed to inhabit mesopelagic
(550–1,000 m deep) waters of tropical and subtropical
Atlantic and Indo-West Pacific regions (Norman 2,000).
The species is thought to live in groups and exhibits diel
vertical migration, based on evidence using sampling with
a closing net (Clarke 1969). During the day, it stays at
550–1,000 m depth and then rises to feed at 100–300 m
depth during the night. Spawning has been suggested to
occur in deep water close to the seafloor (Bruun 1943;
Nesis 1987), a hypothesis supported by the observation of
juveniles (0.5 cm in length) at depths of 1,000–1,750 m
(Clarke 1970)." (Ohkouchi et al., 2013)		 Ohkouchi et al., 2013 95 53 53 N to 42 S (Jereb & Roper, 2005) although "Spirula spirula is a tropical and subtropical ocean species, and its shells have been found on the shores of Canada at 69.75°N (Mercer 1969). This is the same phenomenon as when cuttlebones are washed ashore far from the known range of sepiids (Nesis 1987b): their gas-filled shells float and drift on oceanic currents after the animal has died." (Xavier et al., 2018) (Jereb & Roper, 2005) -42 53 N to 42 S (Jereb & Roper, 2005) (Jereb & Roper, 2005) 4 1 Depth migration: 800–1000 for juveniles and 400-600 in later ontogenetic stages (Lukeneder et al., 2008:178) (Jereb & Roper, 2005; Lukeneder et al., 2008:178; Mutvei, 2017; Ohkouchi et al., 2013; Price et al., 2009; Lukeneder et al., 2008; Warnke et al., 2010) 1 "This is a mesopelagic species, inhabiting from 600 to 700 m during the day and found in depths less than 300 m at night. The capture of young at depths between 1 000 and 1 750 m suggests that females possibly lay eggs on the bottom of continental slopes" (Jereb & Roper, 2005) (Jereb & Roper, 2005; Ohkouchi et al., 2013; Price et al., 2009; Lukeneder et al., 2008; Warnke et al., 2010) 608.8 "the life span is estimated to be about 18 to 20 months." (Jereb & Roper, 2005) (Jereb & Roper, 2005; Price et al., 2009; Lukeneder, 2016; ; Clarke, 1970; Hoffmann et al., 2018) 365.3 "Based on size distribution within a popula-
tion, Clarke (1970) and Nesis (1987) speculated about the life span of
Spirula ranging between 12 and 20 months, which is similar to the
majority of coleoids." (Hoffmann et al., 2018)		 (Jereb & Roper, 2005; Price et al., 2009; Lukeneder, 2016; ; Clarke, 1970; Hoffmann et al., 2018) 456.6 "The species attains sexual maturity at about 30 mm mantle length (after 12 to 15 months of life)," (Jereb & Roper, 2005) (Jereb & Roper, 2005; Price et al., 2009) 365.3 "The species attains sexual maturity at about 30 mm mantle length (after 12 to 15 months of life)," (Jereb & Roper, 2005) (Jereb & Roper, 2005; Price et al., 2009) NA 0 2 1 1 "The ink was more cohesive than observed histioteuthid inks although it did not appear as a pseudomorph" (Lindsay et al. 2020) (Lindsay et al. 2020) 1 single individual observed on slope of Great Barrier Reef, “the Spirula attempted to jet escape five times during the encounter. A partially gas-filled shell would seem to be a hinderance to rapid movement such as jetting, particularly when changing depth (and pressure). Apparently, when rapid escape is needed, jetting is fairly vigorous, though throughout the encounter it appeared that the squid tired…The first time it jetted, it dove about 5 m. The second time was more vigorous, and it dove about 10 m, after which time it remained still for nearly 30 s. It dove again about 5 m the third time, and the fourth diving attempt was weak and the squid did not move far (1 m). During this fourth and weakest attempt at diving, the Spirula also changed trajectory twice (the only time it was observed to do so). It appeared to be tiring out. It remained stationary (again with only the fins beating) for over 2 min before it (presumably) gained enough energy to jet away one last time, inking as it did so. The ink was more cohesive than observed histioteuthid inks although it did not appear as a pseudomorph.” (Lindsay et al. 2020) Lindsay et al. 2020 6 "Spirula mainly feeds on pelagic crustaceans such as copepods, ostracods among others, as evidenced by stomach contents (Kerr, 1931; Nixon and Dilly, 1977; Young, 1977; Nixon and Young, 2003). Observations of sterols support these findings but are also typical for other crustaceans like euphausids, small decapods and mysids that may also represent typical prey (Ballantine et al., 1981). Observed diel migration patterns of Spirula resemble the range of most pelagic crustaceans (Clarke, 1969). Evidence for food sources other than decapods are not available. Recently, Ohkouchi et al. (2013) analysed nitrogen isotopes supporting the idea that Spirula predominantly fed on detritus and zooplankton." (Hoffmann et al., 2018) Ohkouchi et al. 2013
Hoffmann et al., 2018		 0 1 Hoffmann et al., 2018 3 5 "Shells of Spirula were found as stomach contents of terns (Longley, 1930), petrels and albatrosses (Imber, 1973), tuNA(Okutani and Suzuki, 1975; Lansdell and Young, 2007) and the cephalopod Illex argentinus (Santos and Haimovici, 2002)." (Hoffmann et al., 2018) (Hoffmann et al., 2018) (Santos
& Haimovici, 1997) (Imber,
1996) (Okutani
& Suzuki, 1975) (Bourgeois
et al. 2022)		 3 3 "It has not been observed or filmed in its
habitat, but is presumed to inhabit mesopelagic
(550–1,000 m deep) waters of tropical and subtropical
Atlantic and Indo-West Pacific regions (Norman 2,000).
The species is thought to live in groups and exhibits diel
vertical migration, based on evidence using sampling with
a closing net (Clarke 1969). During the day, it stays at
550–1,000 m depth and then rises to feed at 100–300 m
depth during the night. Spawning has been suggested to
occur in deep water close to the seafloor (Bruun 1943;
Nesis 1987), a hypothesis supported by the observation of
juveniles (0.5 cm in length) at depths of 1,000–1,750 m
(Clarke 1970)." (Ohkouchi et al., 2013). "The numbers of specimens in the samples (Tables 1, 2) suggest that Spirula lives in aggregations; in 9 out of 20 samples in which Spirula occur they are in pairs; and four samples, all taken in daylight, contain larger groups of 8-11 individuals." (Clark 1969)		 Ohkouchi et al., 2013 1 "It has not been observed or filmed in its
habitat, but is presumed to inhabit mesopelagic
(550–1,000 m deep) waters of tropical and subtropical
Atlantic and Indo-West Pacific regions (Norman 2,000).
The species is thought to live in groups and exhibits diel
vertical migration, based on evidence using sampling with
a closing net (Clarke 1969). During the day, it stays at
550–1,000 m depth and then rises to feed at 100–300 m
depth during the night. Spawning has been suggested to
occur in deep water close to the seafloor (Bruun 1943;
Nesis 1987), a hypothesis supported by the observation of
juveniles (0.5 cm in length) at depths of 1,000–1,750 m
(Clarke 1970)." (Ohkouchi et al., 2013)		 Ohkouchi et al., 2013 NA 0 NA 69 59 22.6 21 NY
Taonius pavo Taonius pavo 2000 "The species shows an ontogenetic migration to the deeper waters, as the paralarvae live at depths of 300-400 m, juveniles at 600-650 m and adults deeper than 700 m (Lu & Roper, 1979). Moreno et al. (2009) collected eleven paralarvae ranging between "Vertical distribution extends from the upper 200 m for paralarvae to the mid-depths of 600 m for juveniles; ontogenetic descent continues until the mature adults occur at 2 000 m and deeper" (Jereb & Roper, 2010) (Jereb & Roper, 2010; Quetglas et al. 2013) 700 "The species shows an ontogenetic migration to the deeper waters, as the paralarvae live at depths of 300-400 m, juveniles at 600-650 m and adults deeper than 700 m (Lu & Roper, 1979). Moreno et al. (2009) collected eleven paralarvae ranging between 2.5 and 15.6 mm ML within the top 200 m of the water column, between depths of over 800 to 4000 m in the northeast Atlantic. Sampling done between the surface and 1000 m depth in the Pacific Ocean off Oregon, showed that the species was more abundant at 0-500 m than at 0-200 or 0-1000 m (Pearcy, 1965). Abbes (1970) captured two individuals (52 and 72 mm ML) between 1280 and 4000 m depth in the northeast Atlantic and gave a detailed morphological description of the species. Vecchione et al. (2010) reported 23 specimens from 72 to 300 mm ML collected in the northern Mid-Atlantic Ridge." (Quetglas et al. 2013) (Jereb & Roper, 2010; Quetglas et al. 2013) 1 "Taonius pavo was dominant in the midwater tows (n = 236) and 50.6% of all midwater tows contained the species. Three mid-sized specimens (185–215 mm ML) were collected in the bottom tows. Taonius pavo was present in every year sampled, but abundance spiked in 2012, where 117 specimens were collected in 26 tows, with 18 specimens collected in a single tow (PC1205-27). Size ranges were generally similar across cruises, with a size range of 60–300 mm ML, suggesting yearround spawning" Depth Meso- to bathypelagic (Shea et al. 2017) Shea et al. 2017 85 45 45 N to 40 S (Jereb & Roper, 2010) (Jereb & Roper, 2010) -40 "Its main range in the Southern Hemisphere spans 30-40 S" (Imber 1978) (Imber 1978) 3 1 "Vertical distribution extends from the upper 200 m for paralarvae to the mid-depths of 600 m for juveniles; ontogenetic descent continues until the mature adults occur at 2 000 m and deeper" (Jereb & Roper, 2010) (Jereb & Roper, 2010; Shea et al. 2017) NA 0 1 1 1 "Vecchione and Roper (Vecchione and Roper 1991) described ‘J-postures’ and ‘cockatoo postures’ in some deep-sea squid [Taonius pavo, Megalocranchia oceanica, Teuthowenia megalops; Vecchione and Roper 1991. p. 437, 439], and these appear to be other forms of Flamboyant Displays" (Hanlon & Messenger, 2018, p. 126) Vecchione and Roper 1991
Hanlon & Messenger, 2018, p. 12		 0 0 14 "T. pavo appears frequently in the stomach contents of the teuthophagous predators in all world oceans, especially in whales (Yatabe et al., 2010) and also in fur seals (Field et al., 2007), seabirds (Cooper & Klages, 2009), oceanic squids (Watanabe et al., 2004) and deep-sea fishes (Bergstad et al., 2010)" (Quetglas et al. 2013) (Sekiguchi et al. 1996) (Santos
et al. 2007) (Yatabe
et al. 2010) (Mitsui
et al. 2014) (Antonelis
et al. 1987) (Vaske
Júnior et al. 2009) (Quetglas
et al. 2013) (Clarke & Kristensen, 1980) (Chua
et al. 2019) (Fernandez
et al., 2014)		 4 0 not gregarious (inferred from photo and video material) NA 0 NA "Spawning appears to occur suddenly, very near the surface at night." (Jereb & Roper, 2010) 28 25 228.4 540 NY
Teuthowenia megalops Teuthowenia megalops 2700 "The vertical distribution ranges from about 40 m to nearly 2 700 m with evidence of both significant ontogenetic descent as well as some diel vertical movement. Juveniles, subadults and adults inhabit waters where bottom depth exceeds 1 000 m. " (Jereb & Roper, 2010) (Jereb & Roper, 2010; Boyle, 1983; Vecchione & Roper, 1991, p. 437) 40 "The vertical distribution ranges from about 40 m to nearly 2 700 m with evidence of both significant ontogenetic descent as well as some diel vertical movement. Juveniles, subadults and adults inhabit waters where bottom depth exceeds 1 000 m. " (Jereb & Roper, 2010) (Jereb & Roper, 2010; Boyle, 1983; Vecchione & Roper, 1991, p. 437) 1 "lives in epipelagic, mesopelagic and bathypelagic zones, following the
general cranchiid pattern of ontogenetic descent. By full growth, animals have descended into the bathypelagic zone beyond
2 000 m depth, where maturation and mating occur." (Jereb & Roper, 2010)		 Jereb & Roper, 2010 43 74 "Up to 70°N in the Baffin Sea, to 66°N in the Denmark Strait and to 63°N in the Norwegian Sea; recently found in the eastern Greenland Sea about 74°N" (Xavier et al., 2018) (Xavier et al., 2018) 31 31 N to 66 N (Jereb & Roper, 2010) (Jereb & Roper, 2010) 3 1 "The vertical distribution ranges from about 40 m to nearly 2 700 m with evidence of both significant ontogenetic descent as well as some diel vertical movement. Juveniles, subadults and adults inhabit waters where bottom depth exceeds 1 000 m. " (Jereb & Roper, 2010) (Jereb & Roper, 2010) 1 "The vertical distribution ranges from about 40 m to nearly 2 700 m with evidence of both significant ontogenetic descent as well as some diel vertical movement. Juveniles, subadults and adults inhabit waters where bottom depth exceeds 1 000 m. " (Jereb & Roper, 2010) (Jereb & Roper, 2010) 1095.8 2-3 years (Nixon in Boyle (1983), p. 247) (Nixon in Boyle (1983), p. 247) 730.5 2-3 years (Nixon in Boyle (1983), p. 247) (Nixon in Boyle (1983), p. 247) NA 0 3 3 1 "The orange-brown chromatophores are usually retracted to tiny points only increasing in size if the animal is disturbed, but even then it remains largely translucent (Dilly, 1972). The sub-ocular light organs prevent the very large, opaque eyes from revealing the presence of this animal to predators swimming below, by providing a counter-shading mechanism (Fraser, 1962; Clarke, 1963; Denton er al., 1970; Denton, 1971). The other opaque organ, the digestive gland and associated ink sac has a reflective surface (Dilly & Nixon, 1976)" (Boyle, 1983). Boyle, 1983 1 "Vecchione and Roper (Vecchione and Roper 1991) described ‘J-postures’ and ‘cockatoo postures’ in some deep-sea squid [Taonius pavo, Megalocranchia oceanica, Teuthowenia megalops; Vecchione and Roper 1991. p. 437, 439], and these appear to be other forms of Flamboyant Displays" (Hanlon & Messenger, 2018, p. 126) Vecchione and Roper 1991
Hanlon & Messenger, 2018, p. 12		 1 "The curious inking behaviour of the oceanic squid Teuthowenia (formerly Taonius) megalops may fall into the category of protean behaviour. Upon initial disturbance in a shipboard aquarium, small individuals will ink and jet away, but upon further disturbance they begin a ‘balling up’ sequence, in which the fin and eventually most of the mantle and head are inverted into the mantle cavity; the two tentacles are left extended but can be withdrawn too, and the animal will then ink inside the balled-up mantle (Dilly, 1972)." (Hanlon & Messenger, 2018, p. 130) Hanlon & Messenger, 2018, p. 130 "NA" (Xavier et al., 2018) (Xavier et al., 2018) 0 0 10 Clarke and Krstensen (1980) found this species beaks in the stomach of a northern
bottlenosed whale, Hyperoodon ampullatus (Forster, 1770) "Fourteen lower
beaks of this species represent 2-0% of the lower beaks from the Faroes whale
(Table 1). The species contributed 1-2% and 57% of the lower beaks from sperm
whales caught off Iceland and Spain respectively (Clarke & MacLeod,
1974,1976). They comprised over 32% of lower beaks found in blue sharks off
Looe, Cornwall (Clarke & Stevens, 1974). As this is a very common species
in oceanic water north of the latitude of Spain (Lu & Clarke, 1975) it is
not surprising to find it in the diet of predators. All but two of the 14 beaks
have undarkened wings and were therefore from 'immature' squids (Fig. 1)" (Clarke
& Kristensen, 1980) [Species is called Taonius megalops in the article]		 (Jereb
& Roper, 2010) (Fernandez
et al., 2014) (Clarke
& Kristensen, 1980) (Xavier
et al., 2018) (Imber
1978) (Santos
et al. 2006) (Clarke
& Goodall 1994) (Clarke,
1986) (Boyle,
1983).		 4 0 not gregarious (inferred from photo and video material) NA 0 NA 18 15 260.8 265 NY
Todarodes sagittatus Todarodes sagittatus 4595 "it can be found in surface waters above depths of 4595 m (Collins et al., 2001) and as
deep (ROV observation) as 1947 m (Moiseev, 1991). " (Jereb et al., 2015)		 Pierce et al., 2010 0 "They occur in both the open ocean and coastal waters, and from the surface to near‐bottom at depths up to 2500 m." (Pierce et al., 2010) Pierce et al., 2010 2 **edge case** opportunistic sampling over 5-years in NE Atlantic “results indicate that T. sagittatus is a common pelagic and bentho-pelagic species inhabiting deep water (>200 m) to the west of the British Isles particularly in northern areas in late summer and early autumn. All of the samples were caught in demersal trawls which may not be the most effective sampling gear, particularly since dietary analysis indicated that T. sagittatus has a more pelagic habitat.” (Lordan et al. 2001). "lives primarily above the slope and at the bottom near the slope" (Cherel et al., 2009) Lordan et al. 2001; Cherel et al., 2009 90 77 About 77 N to 13 S (Jereb & Roper, 2010) (Jereb & Roper, 2010) (Arkhipkin et al., 2015) (Xavier et al., 2018) -13 "is found throughout the eastern Atlantic to ca. 40°W, and from the Arctic Ocean to ca. 13°S. The range includes the North Sea and the Mediterranean (Clarke, 1966; Zuev et al., 1976; Roper et al., 1984)." (Pierce et al., 2010) (Pierce et al., 2010) (Jereb & Roper, 2010) (Arkhipkin et al., 2015) 2 1 "sagittatus undergoes important trophic and ontogenetic migrations in the North Atlantic Ocean. In early summer large schools appear off the south and southwest coast of Iceland, the Faeroe Islands, Norway and, in some years, Scotland, where they remain until the beginning of winter. Coastal strandings of great numbers of squid are relatively common during this period. As winter arrives, the squid migrate into deeper offshore watersfor the duration of winter. The populations of the northwestern African waters and the western Mediterranean are rather stationary in comparison. This species is found in large numbers from March to May on the fishing grounds around Madeira and other parts of the eastern central Atlantic Ocean. Here ontogenetic movements occur as well, from the shelf to the slope and deep waters." (Jereb & Roper, 2010) (Jereb & Roper, 2010) 1 “One of the main factors affecting growth is temperature but its importance on T. sagittatus would be minimized because, apart from living in a wide depth range both in the Mediterranean and in west Sahara, the squid makes diel vertical movements and also ontogenetic migrations to deeper waters.” (Quetglas & Morales-Nin 2004) (Quetglas & Morales-Nin 2004) (Jereb & Roper, 2010) 1 "sagittatus is known to carry out diel vertical migrations between the surface and near-surface waters at night and in proximity to bottom waters during the day. However, night catches in deeper waters indicate that a portion of the population may not adhere to this general pattern." (Jereb & Roper, 2010) (Jereb & Roper, 2010) (Arkhipkin et al., 2015) 426.125 "Prior to statolith studies, individuals >50 cm were believed to be at least 2 years old (Nesis, 1982/1987), but most recent studies based on statolith increments have sug- gested a longevity of not more than 14 months for individuals reaching up to 47 cm ML (Rosenberg et al., 1981; Wiborg et al., 1982; Arkhipkin et al., 1999; Lordan et al., 2001b; Quetglas and Morales-Nin, 2004; Potoschi et al., 2009)." (Jereb et al, 2015) (Arkhipkin et al., 2015; Jereb et al, 2015) 365.25 "Most individuals probably live 12–14 months, but the lifespan of the largest individuals may approach 2 years." (Jereb et al., 2015) (Arkhipkin et al., 2015; Jereb et al, 2015) 262 "The maximal age of mature females was 262 d (32 cm ML) and 260 d (34.5 cm), that of mature male was 231 d (21 cm ML)"(Nigmatullin et al. 2002) (Arkhipkin et al.1999; Nigmatullin et al. 2002) 170 “The oldest mature female (319 mm ML, 670 g BW, age 262 d) was about a month older than the oldest mature male (201mm ML, 220 g BW, age 231d). The largest specimen (a female of 345 mm ML and 810 g BW) was 260-d old…Males started maturing at ages 170-180 d…At ages 220^230 d, about 75% of males were mature. However, about a ¢fth of males were still immature at these ages, indicating a wide age range of male maturation (Figure 10). Females matured about a month later than males…At ages 4250 d, maturation was rather fast, and about 70% of females became mature” (Arkhipkin et al.1999) (Arkhipkin et al.1999) NA 0 NA 0 58 "The diet of T. sagittatus is composed of fish, crustaceans, and cephalopods; the presence of cannibalism has also been noted. In northern waters, T. sagittatus feeds
primarily on small herring (Clupea harengus) and cod (Gadus morhua; Hernández‐García, 1992; Piatkowski et al., 1998; Quetglas et al., 1999)." (Pierce et al., 2010)		 (Vafidis et al 2008)
Katsanevakis et al 2008
Elsevier, 2014
Jereb et al., 2015
Pierce et al., 2010		 0 0 4 35 (Jereb et al., 2015) "Table 16.3. Prey composition of Todarodes sagittatus, as known from studies in different regions of the eastern Atlantic and the western Mediterranean (compiled from Wiborg et al., 1982 1 , Breiby and Jobling, 1985 2 ; Joy, 1990 3 ; Hernández-García, 1992 4 ; Marabello et al., 1996 5 ; Stowasser, 1997 6 , 2004 7 ; Piatkowski et al., 1998 8 ; Quetglas et al., 1999 9 ; Lordan et al., 2001b 10)."
Taxon Species
Osteichthyes
Acropomatidae Synagrops microlepis (thinlip splitfish) 4
Alepisauridae: Alepisaurus ferox (lancerfish) 10
Ammodytidae: Ammodytes tobianus (sandeel) 1,2 , Ammodytes spp. 6,7
Argentinidae: ArgentiNAsphyraeNA(lesser argentine) 1 , ArgentiNAspp. 10 , Glossanodon leioglossus (smalltoothed argentine) 9
Belonidae: Belone belone (garfish) 10
Caproidae: Capros aper(boarfish) 4,8,
Carangidae: Trachurus trachurus (Atlantic horse mackerel) 10
Centracanthidae: Centracanthus cirrus(curled picarel) 9 , Spicara smaris (picarel) 9
Chauliodontidae: Chauliodus sloani (Sloane's viperfish)5
Clupeidae: Clupea harengus (Atlantic herring) 1,2,10 , Sprattus sprattus 6 , indet. 6,9
Epigonidae: Epigonus telescopus (black cardinal fish) 8
Gadidae: Gadiculus argenteus(silvery pout) 10 , Gadus morhua (Atlantic cod) 2 , Melanogrammus aeglefinus (haddock) 2,6,7 , Merlangius merlangus (whiting) 3,10 , Micromesistius poutassou (blue whiting) 1,2,6,7,10 , Pollachius virens (saithe) 2 , Trisopterus esmarkii (Norway pout) 1,2,3 , Trisopterus minutus (poor cod) 6 , Trisopterus spp. 6,10 , indet. 6,7
Macrouridae: Macrourinae indet. 8 , Nezumia aequalis (common Atlantic grenadier) 9 , indet. 4
Merlucciidae: Merluccius merluccius (European hake) 9
Myctophidae: Benthosema glaciale (glacier lantern fish) 1,9,10 , Ceratoscopelus maderensis (Madeira lanternfish) 9 , Diaphus dumerilii 4 , Diaphus raffinesquii 5 , Diaphus spp. 4,10 , Hygophum benoiti (Benoit’s lanternfish) 5 , Hygophum hygomii (Bermuda lantern fish) 9 , Lampanyctus crocodilus (jewel lanternfish) 5,9 , Myctophum punctatum (spotted lanternfish) 5,10 , Notoscopelus elongatus 9 , Symbolophorus veranyi (large-scale lanternfish) 9 , indet. 4,7,8,10
Moridae: Lepidion lepidion (Mediterranean codling) 9 , Mora moro (common mora) 9
Notacanthidae: Polyacanthonotus rissoanus (smallmouth spiny eel) 9
Osmeridae: Mallotus villosus (capelin) 2
Paralepididae: Arctozenus risso (spotted barracundina) 1,9 , Paralepis spp. 10 , Sudis hyaliNA5 , indet. 5
Phosichthyidae: Ichthyococcus ovatus 5 , Vinciguerria poweriae (Power's deep-water bristle-mouth fish) 5 , Vinciguerria attenuata 5
Scombridae: Scomber scombrus (Atlantic mackerel) 10
Sebastidae: Sebastes spp. 1,2
Soleidae: Microchirus boscanion (Lusitanian sole) 8
Sparidae: Boops boops (bogue) 9
Sternoptychidae: Argyropelecus hemigymnus (half-naked hatchetfish) 5,9,10 , Maurolicus muelleri (pearlside) 1,2,5,7,9,10
Stichaeidae: Leptoclinus maculatus (daubed shanny) 2
Stomiidae: Chauliodus sloani (Sloane's viperfish) 9 , Stomias boa (boa dragonfish) 5,9
Trichiuridae: Lepidopus caudatus (silver scabbardfish) 9
Chondrichthyes
Scyliorhinidae: Galeus melastomus (black-mouthed dogfish) 9 , indet. 9
Crustacea
Decapoda
Dendrobranchiata-Penaeiodea: Aristeus antennatus 9 , Penaidea indet. 4
Macrura reptantia-Astacidea: Nephrops norvegicus (Norway lobster) 9 , indet. 4
Pleocyemata-Anomura: Galatheidae indet. 8 , Munida iris 9 , Munida spp. 4,9
Pleocyemata-Caridea: Alpheus glaber 9 , Crangonidae indet. 9 , Oplophoridae indet. 4 , Pasiphaea multidentata 9 , P. sivado 4,9 , Pasiphaea spp. 2,9,10 , Plesionika giglioli 9 , P. heterocarpus 4,9 , Plesionika spp. 4,8,9 , Processa canaliculata 9 , Processa spp. 9 , indet. 8
Euphausiacea: Meganyctiphanes norvegica 2 , indet. 8,10
Amphipoda: Lyssianassidae indet. 6 , Parathemisto spp. 1,10 , PhrosiNAsemilunata 5 , Phronima spp. 5 , Themisto abyssorum (as Parathemisto abyssorum) 2 , indet. 1,3,10
Isopoda: CirolaNAsp. 5 , Idothea spp. 1 , indet. 1,9
Copepoda: Calanoidea indet. 2 , Pareuchaeta spp. 1 , indet. 1,3,6,7,10
Cephalopoda Indet. 9
Myopsida: Loligo forbesii 9 , Loliginidae indet. 5 , 7
Oegopsida: Abralia veranyi (eye-flash squid) 8 , Abraliopsis spp. 4 , Ancistroteuthis lichtensteini (angel squid) 9 , Brachioteuthis riisei 8 , Cranchiidae indet. 4,8 , Histioteuthis bonnellii (umbrella squid) 5,9 , H. reversa (elongate jewel squid) 9 , Gonatus spp. 1,10 , Illex coindetii (broadtail
shortfin squid) 9,10 , Ommastrephidae indet. 4,6,7,8 , Onychoteuthis banksii (common clubhook squid) 5,9 , Onychoteuthidae indet. 4 , Thysanoteuthis rhombus(diamond squid) 9 , Todarodes sagittatus(European flying squid) 1,2,5,9,10 , Todaropsis eblanae (lesser flying squid) 8 , indet. 8
Sepioidea: Heteroteuthis dispar (odd bobtail) 9 , Neorossia caroli (carol bobtail) 9 , Sepia orbignyaNA4 , Sepia spp. 4,8 , Sepietta neglecta 9 , Sepiola atlantica 3 , Sepiolidae indet. 7,9
Octopoda: Bathypolypus sponsalis (globose octopus) 9 , Eledone cirrhosa (horned octopus) 6 , Octopodidae indet. 6
Gastropoda: indet. 7
Thecosomata: LimaciNAretroversa (retrovert pteropod) 2,10
Bivalvia: indet. 7
Polychaeta: indet. 3
Eunicida: Eunice spp. 2
Phyllodocida: Nereis pelagica (bristlerworm) 2,10 , Nereis spp. 1
Chaetognatha
Sagittoidea: Parasagitta elegans (as Sagitta elegans) 2 , Sagitta spp. 1,2 		(Bloch et al. 2012) (Arkhipkin et al., 2015) (Jereb &
Roper, 2010) (Cherel et al., 2009) (Xavier et al., 2018) (Kousteni et al. 2018)
(Battaglia et al. 2013) (Bello 1998) (Di Lorenzo et al., 2020) (Santos et al.
2006) (Castriota et al. 2015) (Monteiro et al. 2015) (Pierce et al., 2010) (Romeo
et al. 2009) (Peristeraki et al. 2005) (Quetglas et al. 1999) (Jereb et al.,
2015) (Roper & Young 1975).		 4 3 "Foraging shoals of T. sagittatus have been reported from the Arctic since the late 1800s (see Golikov et al., 2013, for detail). Such excursions are described to last for long periods of time and to arise with a certain periodicity. Interestingly, these foraging shoals have not been recorded in Arctic waters between the early 1980s and recent years, apparently reappearing only in 2010 (Golikov et al., 2013). […] From June on, large schools appear off the south and southwest coasts and in the northwestern fjords of Iceland and off the Faroe Islands, Norway, and, in some years, Scotland, where they stay until ca. December (Stephen 1937; Wiborg 1972, 1979a, 1987; Sundet, 1985; Joy, 1990; Boyle et al., 1998; Jonsson, 1998; Lordan et al., 2001b; Bjørke and Gjøsæter, 2004; Roper et al., 2010)" (Jereb et al., 2015) Jereb et al. 2015 1 "Foraging shoals of T. sagittatus have been reported from the Arctic since the late 1800s (see Golikov et al., 2013, for detail). Such excursions are described to last for long periods of time and to arise with a certain periodicity. Interestingly, these foraging shoals have not been recorded in Arctic waters between the early 1980s and recent years, apparently reappearing only in 2010 (Golikov et al., 2013). […] From June on, large schools appear off the south and southwest coasts and in the northwestern fjords of Iceland and off the Faroe Islands, Norway, and, in some years, Scotland, where they stay until ca. December (Stephen 1937; Wiborg 1972, 1979a, 1987; Sundet, 1985; Joy, 1990; Boyle et al., 1998; Jonsson, 1998; Lordan et al., 2001b; Bjørke and Gjøsæter, 2004; Roper et al., 2010)" (Jereb et al., 2015) Jereb et al. 2015 NA 0 NA "The spawning grounds for T. sagittatus are unknown, though this species is thought to spawn extensively throughout the North Atlantic (Shimko 1989; Borges and Wallace 1993). Most immature specimens have been reported from Norwegian waters (Wiborg 1984), and Ireland has been suggested as the closest spawning ground." (Ringvold & Taite 2018) Suggestive “A total of 27 large, gelatinous spherical masses observed in coastal Norwegian waters from Nordland to Aust-Agder Counties in Norway, and off Lysekil in Sweden, Muljica Island in Croatia, Gulf of Naples in Italy, Reqqa Point in Malta, and Saint Mandrier in France, during the months of April to September 2001 to 2017, are reported. Individual spheres measured 0.3–2 m in diameter, averaging one metre (n = 24, ±0.53 m), with all but four sighted in suspension in the water column between 0.5 and 52 m depth…We attribute these gelatinous spheres to the egg masses of squid (Cephalopoda, Oegopsida), and most likely to the ommastrephid Todarodes sagittatus, given similarities with egg masses of T. pacificus…Producing large gelatinous spheres is a good strategy for egg survival…The pressure wave in front of a plankton net easily deflects them, and they are probably hard to eat (O’Dor and Dawe 2013). The transparent jelly balls are difficult to spot, and protect the eggs and larvae from bacteria” (Ringvold & Taite 2018) 149 90 780.624 631.152 290 235 W-W
Todaropsis eblanae Todaropsis eblanae 848 "This squid inhabits mainly the lower sublittoral and upper bathyal throughout the Mediterranean waters (Mangold-Wirz, 1963; Ragonese and Jereb, 1990; Belcari and Sartor, 1993; Salman et al., 1997; Giordano and Carbonara, 1999; Quetglas et al., 2000; Gonzales and Sanchez, 2002; Cuccu et al., 2003; Lefkaditou et al., 2003a, b; Krstulovi  c  Sifner et al., 2005). Deepest records occurred in the northeastern Ionian Sea (848 m; Lefkaditou et al., 2003a), but the species is particularly abundant between 200 and 500 m (e.g., Belcari and Sartor, 1993; Krstulovi  c  Sifner et al., 2005) and in highly productive areas such as the shelf-break (100–200 m; Colloca et al., 2004)." (Arkhipkin et al., 2015) Arkhipkin et al., 2015 20 "in depths between about 20 and 850 m." (Arkhipkin et al., 2015) Arkhipkin et al., 2015 2 **kept as benthic**. JM: maybe should classify as pelagic with rest of Ommastrephidae. "The lesser flying squid is a medium-sized demersal species usually associated with sandy and muddy bottoms," (Arkhipkin et al., 2015). "Todaropsis eblanae (Ball, 1841) is an oceanic squid, which lives near the bottom on the shelf break. In the Mediterranean, it has been observed at depths between 200 and 600 m in the Western area (Quetglas et al., 2000) and in shallower waters in the Central and Eastern areas (Belcari and Sartor, 1993; Tursi and D’Onghia, 1992). In the Atlantic, southeast populations of this species were considered by Roeleveld et al. (1992) as an indicator of the necto-benthic community in the upper slope (300–800 m) off Africa whereas the species occurs on the continental shelf to the west and south of Ireland (Lordan et al., 1998). In spite of a deeper distribution than Loliginids, T. eblanae is said to have lifetraits and behavior more similar to neritic squids than to oceanic ones (Clarke, 1966)." (Robin et al. 2002) Arkhipkin et al., 2015
Lauria et al., 2016
Jereb & Roper, 2010
Pierce et al., 2010
Katsanevakis et al 2008; Robin et al. 2002		 101 61 "The lesser flying squid exhibits a very wide distribution, occurring in shelf waters of the Eastern Atlantic Ocean from 61 N to 36 S, as well as the Baltic Sea and the entire Mediterranean Sea. Its range extends as far north as that of T. sagittatus, beyond the north of Norway (Golikov et al., 2013). However, it also occurs in the western Indian Ocean, western Pacific Ocean, South ChiNASea and Australian waters, the Timor Sea, along the western and eastern Australian coasts, to Tasmania on the eastern side." (Arkhipkin et al., 2015) (Arkhipkin et al., 2015) -40 "The  geographical  distribution  is  discontinuous.  The  species  is  known  from  the 
Mediterranean Sea, the eastern Atlantic from 61°N to 40°S, the southwestern Pacific, 
and the southwestern Indian Ocean off Australia" (Pierce et al., 2010)		 (Katsanevakis et al 2008) (Pierce et al., 2010) 2 0 "The general distribution pattern of T. eblanae in the North Sea as derived from our studies – both in winter and in summer – confirms its preference for the central and northern parts of the North Sea. Todaropsis eblanae is probably the least mobile of the ommastrephid squids in terms of migratory habits, and it tends to behave like a neritic loliginid squid species (Roper et al. 2010)." (Oesterwind et al. 2015) (Oesterwind et al. 2015) 0 "Unlike other ommastrephid species, there is no evidence that T. eblanae regularly as-
cends to the surface or approaches shorelines (Hastie et al., 2009a; Oesterwind et al.,
2010), although it is occasionally caught in coastal waters (Hastie et al., 1994). It is prob-
ably the least mobile of the ommastrephid squids in terms of migratory habits and is
more likely to behave like neritic loliginid squid species than the sympatric om-
mastrephid species I. coindetii and Todarodes sagitattus (Lordan et al., 2001a; Roper et al.,
2010a). " (Jereb et al., 2015)		 (Jereb et al., 2015) 0 "Unlike other ommastrephid species, there is no evidence that T. eblanae regularly as-
cends to the surface or approaches shorelines (Hastie et al., 2009a; Oesterwind et al.,
2010), although it is occasionally caught in coastal waters (Hastie et al., 1994). It is prob-
ably the least mobile of the ommastrephid squids in terms of migratory habits and is
more likely to behave like neritic loliginid squid species than the sympatric om-
mastrephid species I. coindetii and Todarodes sagitattus (Lordan et al., 2001a; Roper et al.,
2010a). " (Jereb et al., 2015)		 (Jereb et al., 2015) 365.25 "The life cycle of T. eblanae is probably annual, because estimated values for the lifespan
range from 7–8 months to 1 year." (Jereb et al., 2015)		 (Arkhipkin et al., 2015; Jereb et al., 2015) 213.0625 "The life cycle of T. eblanae is probably annual, because estimated values for the lifespan
range from 7–8 months to 1 year." (Jereb et al., 2015)		 (Arkhipkin et al., 2015; Jereb et al., 2015) NA 0 NA 0 4 "Like I. coindetii, it is mainly piscivorous, opportunistically on fishes, crustaceans and other cephalopods, in decreasing order of importance; cannibalism also occurs (Rasero et al., 1996)." (Arkhipkin et al., 2015) Arkhipkin et al 2015
Xavier et al 2018
Jereb et al 2015		 0 0 4 26 "Table 17.6. Known predators of Todaropsis eblanae in the Mediterranean Sea and Northeast Atlan-
tic. " (Jereb et al., 2015) Taxon Species References
Cephalopoda Clubhook squid (Onychoteuthis banksii) Hastie et al. (2009a)
Chondrichthyes Black-mouthed dogfish (Galeus melastomus) Kabasakal (2002)
Blue shark (Prionace glauca) Clarke and Stevens (1974)
Portuguese shark (Centroscymnus coelolepis) Hastie et al. (2009a)
Shortfin mako shark (Isurus oxyrinchus) Hastie et al. (2009a)
Sleeper shark (Somniosus spp.) Hastie et al. (2009a)
Smooth hammerhead (SpyrNAzigaena) Hastie et al. (2009a)
Smooth lanternshark (Etmopterus pusillus) Xavier et al. (2012)
Osteichthyes Albacore (Thunnus alalunga) Hastie et al. (2009a)
Atlantic bluefin tuNA(Thunnus thynnus) Karakulak et al. (2009), Romeo et al. (2012)
Swordfish (Xiphias gladius) Hernández-García (1995), Salman (2004), Hastie et al. (2009a)
Aves: Great albatross (Diomedea spp.) Hastie et al. (2009a)
Sooty albatross (Phoebetria fusca) Hastie et al. (2009a)
Cetacea: Bottlenose dolphin (Tursiops truncatus) Santos et al. (1997, 2007), Blanco et al. (2001)
Common dolphin (Delphinus delphis) Pascoe (1986)
Northern bottlenose whale (Hyperoodon ampullatus) Santos et al. (2001c)
Risso’s dolphin (Grampus griseus) Clarke and Pascoe (1985), Bearzi et al. (2011), Bloch et al.
(2012)
Sperm whale (Physeter macrocephalus) Hastie et al. (2009a)
Spotted dolphin (Stenella attenuata) Hastie et al. (2009a)
Striped dolphin (Stenella coeruleoalba) Würtz and Marrale (1993), Blanco et al. (1995)		 (Blanco et al., 2006) (Arkhipkin et al., 2015) (Jereb &
Roper, 2010) (Xavier et al., 2018) (Kousteni et al. 2018) (Di Lorenzo et al.,
2020) (Lipinski & David 1990)  (Velasco
et al., 2001) (Lansdell & Young 2007) (Pierce et al., 2010) (Lansdel and
Young 2007-08) (Jereb et al., 2015) (Varela et al 2018)		 5 0 not gregarious (inferred from photo and video material) NA 0 NA "The spawning grounds are still unknown. However, a possible location was identified
in Northwest Iberian waters, where paralarvae matching the spawning season of T.
eblanae were collected during plankton cruises in many years (Rocha et al., 1999;
Moreno et al., 2009). " (Jereb et al., 2015)		 "No in situ observations on egg deposition are available, although they probably are laid in floating masses, as observed for many other ommastrephids in the natural environment and
in captivity." (Jereb & Roper, 2010)		 "No in situ observations on egg deposition are available, although they probably are laid in floating masses, as observed for many other ommastrephids in the natural environment and
in captivity." (Jereb & Roper, 2010)		 92 63 481.954 334.4 148 106 W-W
Tremoctopus violaceus Tremoctopus violaceus 250 [no source on adults was available so we use juvenile data as an exception] "Observed at the surface at night but may undergo small diel vertical migrations. Juveniles have been collected at depths ranging from 0 to 250 m." (Jereb et al., 2014) (Jereb et al., 2014; Young et al. 1998) 0 [no source on adults was available so we use juvenile data as an exception] "Observed at the surface at night but may undergo small diel vertical migrations. Juveniles have been collected at depths ranging from 0 to 250 m." (Jereb et al., 2014) (Jereb et al., 2014; Young et al. 1998) 1 "epipelagic species" (Laptikhovsky & Salman, 2002) Haimovici et al., 1989;
Laptikhovsky & Salman, 2002		 76 40 40 N to 36 S (Jereb et al., 2014) (Jereb et al., 2014) -36 40 N to 36 S (Jereb et al., 2014) (Jereb et al., 2014) 1 1 "The present is the first Adriatic record of Tremoctopus violaceus since 1936 (Lane, 1974). The find of this pelagic octopod in the Adriatic in late summer may depend on the hydrological properties of the sea. In fact, blanket octopus occurrence in some areas of the Mediterranean seems to be linked to the warming up of the waters (Wirz, 1958; Biagi & Bertozzi, 1992)." (Bello 1993) (Bello 1993; Tubino et al. 2010) 1 "Observed at the surface at night but may undergo small diel vertical migrations. Juveniles have been collected at depths ranging from 0 to 250 m." (Jereb et al., 2014) (Jereb et al., 2014) 730.5 "The life cycle is about 2 years." (Jereb & Roper, 2010) (Jereb & Roper, 2010) 730.5 "The life cycle is about 2 years." (Jereb & Roper, 2010) (Jereb & Roper, 2010) 2 2 1 Tremoctopus violaceus apparently feeds in a similar way, with tactile stimuli to the mantle eliciting arm sweeping movements (Thomas, 1977), but more remarkable is the fact that small individuals of this species, which is often found associated with epipelagic cnidarians, carry fragments of Physalia tentacles in two rows along the first and second pair of arms, corresponding to sucker rows (ibid.]. Jones (1963), who was badly stung when handling Tremoctopus, suggested that the nematocysts could be used mainly as offensive weapons, to capture plankton or small fish that collided with the outspread arms. This remains to be established; it is curious that Physalia fragments are found only in males and in young females" (Hanlon & Messenger, 2018, p. 82) Hanlon & Messenger, 2018, p. 82 1 "For Tremoctopus [violaceus] we could add offence or defence, because the suckers in each row on the dorsal arms of this epipelagic octopod are modified to hold pieces of the Portuguese Man-of-War jellyfish, Physalia, complete with their nematocysts (Jones, 1963)." (Hanlon & Messenger, 2018) Hanlon & Messenger, 2018
Nabhitabhata et al., 2009		 4 3 1 "Males and young females are reported to carry pieces of siphonophore tentacles of the Portuguese Man-of-War Jellyfish (Physalia sp.), by using suckers on their arms I and II. This implies that their function is for defense and catching prey (offense) (Jones 1963, Thomas 1977, Knudsen 1992, Norman et al. 2002). Females with mantle lengths of more than 70 mm have never been reported to function in this way (Thomas 1977)." (Nabhitabhata et al., 2009) Nabhitabhata et al., 2009
Jones 1963		 1 "Ink sac present." (Jereb et al., 2014) (Jereb et al., 2014) 1 the octopus became threatened, extended her web, and jettisoned her eggs" (Jiménez-Badillo et al. 2021) Jiménez-Badillo et al. 2021 1 "In case of danger, distal segments of the arms with their large lateral web are detached " (Orsi Relini 2009) Orsi Relini 2009 4 "Female Tremoctopus violaceus are reported to feed on pteropod molluscs and small fishes." (Jereb et al., 2014) Jereb et al 2014 0 0 4 17 "Vaske-Junior
& Rincón-Filho (1998) reported the presence of T. violaceus in the stomach
contents of blue sharks (Prionace glauca) caught by longline fishing off the
coast of Rio Grande do Sul and Santa CatariNAstates (southern Brazil). VaskeJunior
(2005) recorded beaks and semi-digested individuals of T. violaceus in stomach
contents of yellowfin tuNA(Thunnus albacares), Atlantic albacore (T.
alalunga), Bigeye tuNA(T. obesus), Wahoo (Acanthocybium solandri), common
dolphinfish (CoryphaeNAhippurus), swordfish (Xiphias gladius), Atlantic white
marlin (Tetrapturus albidus), Longbill spearfish (T. pfluegeri), black marlin,
(Makaira nigricans), Atlantic sailfish (Istiophorus albicans), Longnose
lancetfish (Alepisaurus ferox), blue shark (Prionace glauca), Night shark
(Carcharhinus signatus) and Scalloped hammerhead (SphyrNAlewini), catched in
Northeastern Brazil." (Tubino
et al. 2010)		 (Lopes et al. 2012) (Tubino et al. 2010) (Vaske Júnior et
al. 2009) (Tscuchiya et al. 1998) (Bello 1993) (Salman & Karakulak 2009) (Castriota
et al 2008) (Battaglia et al. 2013)		 3 following from FAO Jereb et al. 2014 "Known to occur, on occasion, in plague proportions" but no further specification or reference (Jereb et al., 2014) 0 not gregarious (inferred from photo and video material) 2 1 1 "For Tremoctopus [violaceus] we could add offence or defence, because the suckers in each row on the dorsal arms of this epipelagic octopod are modified to hold pieces of the Portuguese Man-of-War jellyfish, Physalia, complete with their nematocysts (Jones, 1963)." (Hanlon & Messenger, 2018) Jones, 1963 in Hanlon & Messenger 2018 1 1 "Females brood eggs within the arm crown and webs, attached as strings to a secreted, mineralized rod. Egg strings at different stages of development indicate prolonged spawning and potentially high fecundity. Up to five different egg stages can be carried by a single female." (Jereb et al., 2014) Jereb et al., 2014 0 "Females brood eggs within the arm crown and webs, attached as strings to a secreted, mineralized rod. Egg strings at different stages of development indicate prolonged spawning and potentially high fecundity. Up to five different egg stages can be carried by a single female." (Jereb et al., 2014) Jereb et al., 2014 56 23 332.008 158.4 24.44 71 15 W-NY-W The smallest estimate is for a male. It's quite a bit smaller than the other two estimates, due to the extreme sexual dimorphism in this species.
Vampyroteuthis infernalis Vampyroteuthis infernalis 3300 "The vertical distribution of V infernalis
covers meso- and bathypelagic zones from 600 to 3300 m depth" (Golikov et al., 2019)		 (Cairns, 1976; Shea et al. 2017; Timme et al., 2020; Judkins & Vecchione, 2020; Jereb et al., 2014; Golikov et al., 2019; Hoving et al., 2015; Bower et al., 2006; Robison et al., 2003; Robison et al., 2003; Young, 1972; Norman and Reid 2000 – Book, p19; Norman and Reid 2000 – Book, p50; Vecchione & Roper, 1991, p. 437; Roper & Young 1975; Roper & Young 1975; Seibel et al., 1997; Hoving & Robison, 2012) 200 "Among 157 Vampyroteuthis infernalis individuals, the majority
were below 600 m both day and night (4–135 mm ML). Two
individuals (ML = 5 mm, 7 mm) were found between 200 and
600 m. We do not believe that this depth layer is characteristic of juveniles as over 30 individuals of similar MLs found between 600 and 1500 m. Vampires are non-migrators and do not appear to exhibit an ontogenic shift during their life history" (Judkins & Vecchione, 2020)		 (Cairns, 1976; Shea et al. 2017; Timme et al., 2020; Judkins & Vecchione, 2020; Jereb et al., 2014; Golikov et al., 2019; Hoving et al., 2015; Bower et al., 2006; Robison et al., 2003; Robison et al., 2003; Young, 1972; Norman and Reid 2000 – Book, p19; Norman and Reid 2000 – Book, p50; Vecchione & Roper, 1991, p. 437; Roper & Young 1975; Roper & Young 1975; Seibel et al., 1997; Hoving & Robison, 2012) 1 "a mid-water species" (Jereb et al., 2014) Jereb et al., 2014 96 61 "Our specimen from the Mid-Atlantic Ridge (61.5°N, 30.4°W; 11 – 1; Supplementary Table 1) represents the current northernmost record ofthe species. " (Golikov et al., 2019) (Golikov et al., 2019) -35 "between approximately 35°N and 35°S. " (Jereb et al., 2014) (Jereb et al., 2014) 4 0 "Vampires are non-migrators and do not appear to exhibit an ontogenic shift during their life history" (Judkins & Vecchione, 2020) (Judkins & Vecchione, 2020) 0 "A significant ontogenetic increase of 𝛿^13C in V infernalis, coupled with lack of significant depth-related trends, suggests that there are no ontogenetic depth preferences in V infernalis." (Golikov et al., 2019) (Golikov et al., 2019) 2922 "If we assume 38–100 spawning events (as in our most advanced female), the duration of the adult stage is at least 3–8 years in this specimen, with the total lifespan exceeding these numbers" (Hoving et al., 2015) (Hoving et al., 2015) 1095.8 "If we assume 38–100 spawning events (as in our most advanced female), the duration of the adult stage is at least 3–8 years in this specimen, with the total lifespan exceeding these numbers" (Hoving et al., 2015) (Hoving et al., 2015) 2 2 1 "they possess several bioluminescent displays, which are believed to be incorporated into anti-predation behaviour and perhaps to prey capture" (Hoving & Robison, 2012) Hoving & Robison, 2012 1 "In aquarium experiments, Hunt [4] presented live Artemia nauplii to Vampyroteuthis with extended filaments. When the nauplii contacted the filament, the vampire squid swam around the location where the nauplii touched the filament and enveloped them within its webbed arms." (Hoving & Robison, 2012) Hoving & Robison, 2012
Hunt 1996		 5 5 1 "they possess several bioluminescent displays, which are believed to be incorporated into anti-predation behaviour and perhaps to prey capture" (Hoving & Robison, 2012) Hoving & Robison, 2012
Robison et al., 2003
Herring et al., 1994		 1 "they possess several bioluminescent displays, which are believed to be incorporated into anti-predation behaviour and perhaps to prey capture" (Hoving & Robison, 2012) Hoving & Robison, 2012 1 "Disturbed animals pull the arms and webs over their body to take on an inverted shape that exposes the black skin and cirri, on the oral surfaces of the webs." (Jereb et al., 2014) Jereb et al., 2014
Robison et al., 2003		 0 no ink sac Robison
et al. 2003; Bush & Robison, 2007		 0 "The archaic, bathypelagic cephalopod Vampyroteuthis infernalis releases a
cloud of dimly glowing particles from its arm tips (Robison
et al. 2003), but it has no ink sac." (Bush & Robison, 2007)		 (Robison
et al. 2003; Bush & Robison, 2007)		 1 "Seibel et al. [45] estimate that only short duration burst swimming is avail- able or necessary for predator avoidance, because cryptic coloration and bioluminescent countermeasures [15] prob- ably contribute more to survival." (Hoving & Robison, 2012) Hoving & Robison, 2012
Seibel et al., 1998		 1 "Ann-tip light organs, which can be bitten or broken off and then regenerated, may serve as sacrificial diversions for predators (Herring, 1977). Tip lights are found in several deep-living squids such as Chiroteuthis and Octopoteuthis, where they may also serve as lures for prey, thus functioning like the escae of anglerfish and the barbels of stomiid fishes (Herring, 1977; Young, 1983). Our observations of apparently regenerated light organs at the ends of shortened arms in Vampyroteuthis may be evidence of their potential as sacrificial structures. The characteristics of the arm-tip displays indicate that there is direct neural control of their luminescence." (Robison et al., 2003) Robison et al., 2003 19 "Vampire squid
feed opportunistically on zooplankton
and detritus " (Hoving et al., 2015)		 Golikov et al., 2019
Hoving et al 2015
Schawrz et al 2020
Hoving & Robison, 2012 		0 1 Golikov et al., 2019
Hoving & Robison, 2012		 5 5 Found
in the stomach of the blue shark (Prionace glauca) (Markaida &
Sosa-Nishizaki, 2010) and Brazilian coast (Vaske Júnior et al. 2009) and Ivory
Coast (Konan et al. 2018)		 (Konan et al. 2018) (Fernandez et al., 2014) (Cherel et al.,
2009) (Robison et al., 2003) (Clarke & Kristensen, 1980) (Antonelis et al.
1987) (Sekiguchi et al. 1996) (Imber, 1996)		 2 0 not gregarious (inferred from photo and video material) NA 0 NA [Not necessarily communication, but the only information available] "thout the tip lights glowing as well. On one occasion, male and female specimens were collected on the same day and were then placed in separate kreisels less than a meter apart, in the darkened laboratory ashore. When the female was disturbed and began to flash her arm-tip lights, the undisturbed male quickly and vigorously responded with tip-light flashes. This reaction was repeated twice (Hunt, 1996). We saw no evidence of differential light production by females and males." (Robison et al., 2003) 58 37 209.6 47 45 45 C22-NY
Vitreledonella richardi Vitreledonella richardi 2005 "2005 m" (Sajikumar et al., 2016) (Jereb et al., 2014; Sajikumar et al., 2016; Land 1992; Norman and Reid 2000 – Book, p53; Clarke, 1986; Vecchione & Roper, 1991, p. 438) 50 "…50-500 m depth to deeper than 1000 m…" (Clarke, 1986). (Jereb et al., 2014; Sajikumar et al., 2016; Land 1992; Norman and Reid 2000 – Book, p53; Clarke, 1986; Vecchione & Roper, 1991, p. 438) 1 "Depth range from near the surface to at least 1 000 m, typically over deep water (beyond continental shelf)" (Jereb et al., 2014) Jereb et al., 2014 130 70 70 N to 60 S (Jereb et al., 2014) (Jereb et al., 2014) -60 70 N to 60 S (Jereb et al., 2014) (Jereb et al., 2014) 4 1 possible ontogenetic "Small specimens of less than 20mm ML are found
between 50 and 300 m, and larger animals in depths of 100-l000m
[Clarke and Lu 1974, Land 1992']." (Nixon & Young 2003)		 (Nixon & Young 2003) NA 0 2 1 1 “When the animal is oriented with its body axis 45° from the horizontal the longitudinal axes of the eyes and of the digestive gland are parallel, and vertical. This has the effect of reducing the size of the opaque parts of the animal when it is viewed from below in silhouette; this camouflage strategy is seen in other midwater animals (Land 1992}”(Nixon & Young 2003) (Nixon & Young 2003 1 (Jereb et al., 2014) 0 0 0 not gregarious (inferred from photo and video material) 1 0 1 1 "the female is thought to brood her eggs
in the space formed when her arms are held together." (Norman and Reid 2000 – Book, p53)		 Norman and Reid 2000 – Book, p53) 0 "the female is thought to brood her eggs
in the space formed when her arms are held together." (Norman and Reid 2000 – Book, p53)		 Norman and Reid 2000 – Book, p53) 6 3 16.6 66 NY

Phylogeny

brain_taxa
Superorder
Order
Family
sister on current tree
source
substitute species on genetic phylogeny
Alloteuthis_media Decapodiformes Myopsida Loliginidae Alloteuthis_subulata anderson2000
Alloteuthis_subulata Decapodiformes Myopsida Loliginidae Alloteuthis_media anderson2000
Loligo_forbesii Decapodiformes Myopsida Loliginidae Loligo_vulgaris lindgren2012
Loligo_vulgaris Decapodiformes Myopsida Loliginidae Loligo_forbesii lindgren2012
Lolliguncula_brevis Decapodiformes Myopsida Loliginidae Pickfordiateuthis_pulchella andersonmarian2020
Sepioteuthis_sepioidea Decapodiformes Myopsida Loliginidae Sepioteuthis_lessoniana sales2013, anderson2000
Sepioteuthis_lessoniana Decapodiformes Myopsida Loliginidae Sepioteuthis_sepioidea sales2013
Pickfordiateuthis_pulchella Decapodiformes Myopsida Loliginidae Lolliguncula_brevis andersonmarian2020
Architeuthis_dux Decapodiformes Oegopsida Architeuthidae Neoteuthis_thielei lindgren2012, fernández-alvarez2022
Chiroteuthis_veranii Decapodiformes Oegopsida Chiroteuthidae Grimalditeuthis_bonplandi lindgren2012, fernández-alvarez2022
Grimalditeuthis_bonplandi Decapodiformes Oegopsida Chiroteuthidae Chiroteuthis_veranyi lindgren2012, fernández-alvarez2022
Cranchia_scabra Decapodiformes Oegopsida Cranchiidae Leachia_dislocata lindgren2012
Teuthowenia_megalops Decapodiformes Oegopsida Cranchiidae Taonius_pavo lindgren2012
Bathothauma_lyromma Decapodiformes Oegopsida Cranchiidae Teuthowenia_megalops fernández-alvarez2022
Egea_inermis Decapodiformes Oegopsida Cranchiidae Megalocranchia_maxima sanchez2018, fernández-alvarez2022
Galiteuthis_glacialis Decapodiformes Oegopsida Cranchiidae Helicocranchia_papillata sanchez2018 Galiteuthis_armata
Helicocranchia_papillata Decapodiformes Oegopsida Cranchiidae Galiteuthis_glacialis sanchez2018 Helicocranchia_pfefferi
Leachia_dislocata Decapodiformes Oegopsida Cranchiidae Cranchia_scabra sanchez2018 Leachia_atlantica
Megalocranchia_maxima Decapodiformes Oegopsida Cranchiidae Egea_inermis sanchez2018, fernández-alvarez2022 Megalocranchia_oceanica
Sandalops_melancholicus Decapodiformes Oegopsida Cranchiidae Taonius_pavo evans2018; judkins2022
Taonius_pavo Decapodiformes Oegopsida Cranchiidae Sandalops_melancholicus lindgren2012
Discoteuthis_laciniosa Decapodiformes Oegopsida Cycloteuthidae (Abraliopsis_morisii…Chiroteuthis_veranyi) fernández-alvarez2022
Abraliopsis_morisii Decapodiformes Oegopsida Enoploteuthidae Abralia_veranyi fernández-alvarez2022
Abralia_veranyi Decapodiformes Oegopsida Enoploteuthidae Abraliopsis_morisii fernández-alvarez2022
Gonatus_fabricii Decapodiformes Oegopsida Gonatidae Onychoteuthis_banksii lindgren2012
Histioteuthis_miranda Decapodiformes Oegopsida Histioteuthidae Architeuthis_dux…Onychoteuthis_banksii) lindgren2012
Joubiniteuthis_portieri Decapodiformes Oegopsida Joubiniteuthidae Mastigoteuthis_schmidti lindgren2012, sanchez2018, fernández-alvarez2022
Lycoteuthis_lorigera Decapodiformes Oegopsida Lycoteuthidae Pyroteuthis_margaritifera fernández-alvarez2022
Mastigoteuthis_schmidti Decapodiformes Oegopsida Mastigoteuthidae Joubiniteuthis_portieri lindgren2012, sanchez2018, fernández-alvarez2022 Mastigoteuthis_agassizii
Neoteuthis_thielei Decapodiformes Oegopsida Neoteuthidae Architeuthis_dux lindgren2012, fernández-alvarez2022
Octopoteuthis_danae Decapodiformes Oegopsida Octopoteuthidae (Chiroteuthis_veranyi…Joubiniteuthis_portieri) fernández-alvarez2022
Illex_illecebrosus Decapodiformes Oegopsida Ommastrephidae Illex_coindetii fernández-alvarez2022
Illex_coindetii Decapodiformes Oegopsida Ommastrephidae Illex_illecebrosus fernández-alvarez2022
Todaropsis_eblanae Decapodiformes Oegopsida Ommastrephidae (Illex_illecebrosus,Illex_coindetii) fernández-alvarez2022
Todarodes_sagittatus Decapodiformes Oegopsida Ommastrephidae (Illex_illecebrosus,Illex_coindetii), Todaropsis_eblanae fernández-alvarez2022
Onychoteuthis_banksii Decapodiformes Oegopsida Onychoteuthidae Gonatus_fabricii lindgren2012
Ancistroteuthis_lichtensteinii Decapodiformes Oegopsida Onychoteuthidae Histioteuthis_miranda fernández-alvarez2022
Pterygioteuthis_giardi_hoylei Decapodiformes Oegopsida Pyroteuthidae (Gonatus_fabricii, Onychoteuthis_banksii) lindgren2012
Pyroteuthis_margaritifera Decapodiformes Oegopsida Pyroteuthidae Lycoteuthis_lorigera fernández-alvarez2022
Idiosepius_paradoxus Decapodiformes Idiosepida Idiosepiidae (Neorossia…Loligo) andersonlindgren2021
Sepia_officinalis Decapodiformes Sepiida Sepiidae Sepia_orbignyana, Sepia_elegans lindgren2012
Sepia_bandensis Decapodiformes Sepiida Sepiidae Sepia_plangon lupse2023
Sepia_plangon Decapodiformes Sepiida Sepiidae Sepia_bandensis lupse2023
Sepia_elegans Decapodiformes Sepiida Sepiidae Sepia_orbignyana lupse2023
Sepia_orbignyana Decapodiformes Sepiida Sepiidae Sepia_elegans lupse2023
Neorossia_caroli Decapodiformes Sepiolida Sepiolidae Rossia_macrosoma sanchez2021
Sepietta_oweniana Decapodiformes Sepiolida Sepiolidae Sepietta_obscura sanchez2021
Sepiola_rondeleti Decapodiformes Sepiolida Sepiolidae (Sepietta_obscura, Sepietta_oweniana) sanchez2021
Sepiola_affinis Decapodiformes Sepiolida Sepiolidae Sepiola_robusta, Sepiola_rondeleti sanchez2021
Sepiola_robusta Decapodiformes Sepiolida Sepiolidae Sepiola_rondeleti, Sepiola_affinis sanchez2021
Rossia_macrosoma Decapodiformes Sepiolida Sepiolidae Neorossia_caroli sanchez2021
Heteroteuthis_dispar Decapodiformes Sepiolida Sepiolidae (Neorossia_caroli, Rossia_macrosoma) sanchez2021
Spirula_spirula Decapodiformes Spirulida Spirulidae Bathyteuthida + Oegopsida lindgren2012
Bathyteuthis_abyssicola Decapodiformes Bathyteuthida Bathyteuthidae Chtenopteryx_sicula lindgren2012
Chtenopteryx_sicula Decapodiformes Bathyteuthida Chtenopterygidae Bathyteuthis_abyssicola lindgren2012
Vampyroteuthis_infernalis Octopodiformes Vampyromorpha Vampyroteuthidae Cirrothauma_murrayi…Haliphron_atlanticus lindgren2012
Cirroteuthis_muelleri Octopodiformes Octopoda (Cirrata) Cirroteuthidae Cirrothauma_murrayi sanchez2018
Cirrothauma_murrayi Octopodiformes Octopoda (Cirrata) Cirroteuthidae Cirroteuthis_muelleri lindgren2012
Haliphron_atlanticus Octopodiformes Octopoda (Incirrata) Alloposidae Tremoctopus_violaceus lindgren2012
Tremoctopus_violaceus Octopodiformes Octopoda (Incirrata) Tremoctopodidae Haliphron_atlanticus lindgren2012
Bolitaena_pygmaea Octopodiformes Octopoda (Incirrata) Amphitretidae/Bolitaenidae Japetella_diaphana lindgren2012
Japetella_diaphana Octopodiformes Octopoda (Incirrata) Amphitretidae/Bolitaenidae Bolitaena_pygmaea lindgren2012
Vitreledonella_richardi Octopodiformes Octopoda (Incirrata) Amphitretidae Amphitretus_pelagicus lindgren2012
Amphitretus_pelagicus Octopodiformes Octopoda (Incirrata) Amphitretidae Vitreledonella_richardi sanchez2018
Argonauta_argo Octopodiformes Octopoda (Incirrata) Argonautidae Ocythoe_tuberculata sanchez2018
Ocythoe_tuberculata Octopodiformes Octopoda (Incirrata) Ocythoidae Argonauta_argo sanchez2018
Bathypolypus_sponsalis Octopodiformes Octopoda (Incirrata) Bathypolypodidae Bathypolypus_bairdii strugnell2013
Bathypolypus_bairdii Octopodiformes Octopoda (Incirrata) Bathypolypodidae Bathypolypus_sponsalis strugnell2013
Eledone_cirrhosa Octopodiformes Octopoda (Incirrata) Eledonidae Eledone_moschata lindgren2012
Eledone_moschata Octopodiformes Octopoda (Incirrata) Eledonidae Eledone_cirrhosa lindgren2012
Enteroctopus_dofleini Octopodiformes Octopoda (Incirrata) Enteroctopodidae Argonauta…Scaergus lindgren2012
Callistoctopus_macropus Octopodiformes Octopoda (Incirrata) Octopodidae Macrotritopus_defilippi, Scaeurgus_unicirrhus strugnell2013
Hapalochlaena_fasciata Octopodiformes Octopoda (Incirrata) Octopodidae Abdopus_capricornicus, (Octopus_bimaculatus…vulgaris) strugnell2013
Macrotritopus_defilippi Octopodiformes Octopoda (Incirrata) Octopodidae Callistoctopus_macropus, Scaeurgus_unicirrhus strugnell2013
Octopus_cyanea Octopodiformes Octopoda (Incirrata) Octopodidae (Octopus_vulgaris, Octopus_salutii, Octopus_bimaculatus) taite2023
Octopus_bimaculatus Octopodiformes Octopoda (Incirrata) Octopodidae Octopus_vulgaris, Octopus_salutii taite2023; sánchez-márquez2023
Octopus_vulgaris Octopodiformes Octopoda (Incirrata) Octopodidae Octopus_salutii, Octopus_bimaculatus taite2023; sánchez-márquez2023
Scaeurgus_unicirrhus Octopodiformes Octopoda (Incirrata) Octopodidae Callistoctopus_macropus, Macrotritopus_defilippi strugnell2013
Octopus_salutii Octopodiformes Octopoda (Incirrata) Octopodidae Octopus_vulgaris, Octopus_bimaculatus sánchez-márquez2023
Pteroctopus_tetracirrhus Octopodiformes Octopoda (Incirrata) Octopodidae Scaeurgus_unicirrhus taite2023; sánchez-márquez2023
Abdopus_capricornicus Octopodiformes Octopoda (Incirrata) Octopodidae Hapalochlaena…Octopus strugnell2013 Abdopus_aculeatus

For more details, see:

Basava, K., Bendixen, T., Leonhard, A., George, N. L., Vanhersecke, Z., Omotosho, J., Mather, J. & Muthukrishna, M. (2024). Ecological not social factors explain brain size in cephalopods. https://www.biorxiv.org/content/10.1101/2024.05.01.592020v3

Basava, K., Bendixen, T., Birk Sorensen, A. L., George, N. L., Vanhersecke, Z., Omotosho, J., Mather, J. & Muthukrishna, M. (2024). A phylogeny of extant coleoid cephalopods with brain data. https://www.biorxiv.org/content/10.1101/2024.04.29.591691v2