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Cephalopod size

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Body variation
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Thegiant squid (Architeuthis dux, pictured) was for a long time thought to be the largest extant cephalopod. It is now known that thecolossal squid (Mesonychoteuthis hamiltoni) attains an even greater maximum size. The giant squid seen here measured 9.24 m (30.3 ft) in total length and had amantle length of 1.79 m (5.9 ft).

Cephalopods, which include squids and octopuses, vary enormously in size. The smallest are only about 1 centimetre (0.39 in) long and weigh less than 1 gram (0.035 oz) at maturity, while thegiant squid can exceed 10 metres (33 ft) in length and thecolossal squid weighs close to half a tonne (1,100 lb), making them the largest livinginvertebrates. Living species range in mass more than three-billion-fold,[nb 1] or across nineorders of magnitude, from the lightest hatchlings to the heaviest adults.[4] Certain cephalopod species are also noted for havingindividual body parts of exceptional size.[5]

Cephalopods were at one time thelargest of all organisms on Earth,[6] and numerous species of comparable size to the largest present day squids are known from thefossil record, including enormous examples ofammonoids,belemnoids,nautiloids,orthoceratoids,teuthids, andvampyromorphids. In terms of mass, the largest of all known cephalopods were likely the giant shelled ammonoids andendocerid nautiloids,[7] though perhaps still second to the largest living cephalopods when consideringtissue mass alone.[8]

Cephalopods vastly larger than either giant or colossal squids have been postulated at various times. One of these was theSt. Augustine Monster, a large carcass weighing several tonnes that washed ashore on theUnited States coast nearSt. Augustine,Florida, in 1896. Reanalyses in 1995 and 2004 of the original tissue samples—together with those ofother similar carcasses—showed conclusively that they were all masses of thecollagenous matrix of whaleblubber.[9]

Giant cephalopods have fascinated humankind for ages. The earliest surviving records are perhaps those ofAristotle andPliny the Elder, both of whom described squids of very large size.[10] Tales of giant squid have been common among mariners since ancient times, and may have inspired the monstrouskraken ofNordic legend, said to be as large as an island and capable of engulfing and sinking any ship.[11] Similar tentacledsea monsters are known from other parts of the globe, including theAkkorokamui ofJapan andTe Wheke-a-Muturangi ofNew Zealand. TheLusca of theCaribbean andScylla inGreek mythology may also derive from giant squid sightings,[12] as might eyewitness accounts of other sea monsters such assea serpents.[13]

Cephalopods of enormous sizehave featured prominently in fiction.[14] Some of the best known examples include the giant squid fromJules Verne's 1870 novelTwenty Thousand Leagues Under the Seas andits various film adaptations; the giant octopus from the 1955monster movieIt Came from Beneath the Sea; and the giant squid fromPeter Benchley's 1991 novelBeast and theTV film adaptation of the same name.

Due to its status as acharismatic megafaunal species, the giant squid has been proposed as an emblematic animal formarine invertebrate conservation.[15] Life-sized models of the giant squid are a common sight innatural history museums around the world,[16] and preserved specimens are much sought after for display.[17]

Size in teuthology

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Standard measurements for cephalopods with a squid and octopus as examples

Mantle length

[edit]
Scientists measuring the mantle width of a large female giant squid ofc. 2 m (6.6 ft) ML

Mantle length (ML) is the standard size measure forcoleoid cephalopods (shell diameter being more common fornautiluses) and is almost universally reported in the scientific literature. Themantle is the cephalopod's "body", lying posterior to the head and enclosing thevisceral mass andmantle cavity, the latter being used for locomotion byjet propulsion. Unless otherwise indicated, mantle length is measureddorsally over the midline of the mantle (sometimes specified asdorsal mantle length, DML). It is a straight-line measure, not measured over the curve of the body. InDecapodiformes (ten-limbed cephalopods), mantle length is measured from the anterior edge of the mantle (near the head), to the posterior end of the mantle or the apex of the unitedfins, whichever is longer. InOctopodiformes (eight-limbed cephalopods), the anterior edge of the mantle is not clearly delimited dorsally due to advanced head–mantle fusion, and mantle length is therefore taken from the midpoint between the eyes to the posterior end of the mantle. Whenventral mantle length is meant instead of dorsal this is always specified as such and abbreviated VML.[18]

As an indication of overall size, mantle length is generally considered more reliable than total length becausecephalopod limbs may easily be stretched beyond their natural length and are often damaged or missing in preserved specimens (this is particularly true of the longtentacles of many squid species).[19] Nevertheless, mantle length is not equally applicable to all species. Certainbenthic octopuses such asCallistoctopus ornatus are able to elongate and retract their mantles and therefore mantle length measurements, even when taken from a live specimen, may vary considerably. Another problematic case is that of the gelatinouscirroteuthids, whose weakly muscled mantles are prone to substantial shrinkage during preservation. The interocular distance may be a more reliable standard for this group.[20]

Total length

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Total length (TL) is measured along thedorsal midline with thelimbs outstretched and in line with the body axis. It is the greatest measurable extent of a specimen from the posterior end of the mantle orfins (ortail, if present[21]) to the apex of the longest limb.[18] It is recommended that arms and tentacles be measured in a relaxed state so as not to exaggerate their length, but historically this practice was not always followed and some of the more extreme publishedgiant squid measurements have been attributed to artificial lengthening of the tentacles.[22] Although total length is often mentioned in relation to the largest cephalopod species, it is otherwise seldom used in teuthology.[23] As with mantle length, it is a straight-line measure.

Total length is not to be confused with arm span—also known asarm spread,radial span, orradial spread—which may be much larger and is often reported for octopuses (for which the arms usually constitute the vast majority of the length). In squids, total length is inclusive of the feeding tentacles, which in some species may be longer than the mantle, head, and arms combined (chiroteuthids such asAsperoteuthis acanthoderma being a prime example).

A related measure is standard length (SL), which is the combined length of the mantle, head, and arms, excluding the often long feeding tentacles.[24] This measure is particularly useful for species such as the giant squid, where almost the entire bulk of the animal takes up less than half of its total length.

Mass

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Mass (often abbreviated WT for 'weight') is reported far less frequently than either mantle or total length, and accurate records do not exist for all of the large cephalopod species. It can also vary widely depending on the state of the specimen at the time of weighing (for example, whether it was measured live or dead, wet or dry, frozen or thawed, pre- or post-fixation, with or without egg mass, and so on).

Giant squid beak and associated muscles with hand for scale

Methods of size determination

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In contrast to the vast majority of living cephalopods, which are wholly soft-bodied, size determination of thefew surviving shelled species (in terms of shell diameter) is comparatively straightforward and can be accomplished with a high level of precision. Whatever the type of cephalopod, in the absence of whole specimens, size can often be estimated from only partial remains. For example,cephalopod beaks can be used for mantle length, total length and body mass estimation,[25] and this method has notably been used to estimate the maximum size of thecolossal squid. Thelower rostral length (LRL) of the beak is often used for this purpose. The rostral length of the lower and upper beaks is the standard measure of beak size inDecapodiformes;hood length is preferred forOctopodiformes.[26]

Mantle length has been estimated from video recordings of squid in the wild.[27]

Early life stages

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Hatchlings

[edit]

Hatchlings ofIdiosepius thailandicus, possibly the smallest extant cephalopod species at maturity, have amantle length of around 1 mm (0.039 in).[28] The closely relatedIdiosepius pygmaeus weighs only 0.00033 g (1.2×10−5 oz) upon hatching and increases in weight to 0.175 g (0.0062 oz) as it reaches maturity in 50 days.[2] Even smaller are the hatchlings of the commercially importantIllex illecebrosus, with a mass of 0.00015 g (5.3×10−6 oz).[29] Hatchlings of thegiant Pacific octopus (Enteroctopus dofleini)—one of the two largest octopus species—weigh 0.0253 g (0.00089 oz) on average.[30]

At the other extreme arenautiluses, which upon hatching typically have a shell diameter of 25 mm (1 in) or more (depending on the species), the largest hatchling size among extant invertebrates.[31] Hatchlings ofNautilus belauensis, one of the larger species, are estimated to weigh on the order of 5.9 g (0.21 oz)[nb 2] and mature at around 1.2 kg (2.6 lb) after almost 4000 days, or around 11 years.[2]

Smallest adult size

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Idiosepius paradoxus, a tiny species of squid that grows to 16 mm (0.63 in) in mantle length
Adult maleTremoctopus violaceus with largehectocotylus (arm modified forspermatophore transfer)

The smallest adult size among living cephalopods is attained by the so-called pygmy squids,Idiosepius,[32] and certain diminutive species of the genusOctopus, both of which weigh less than 1 gram (0.035 oz) at maturity.[33]Idiosepius thailandicus is perhaps the smallest of all, with females averaging 10.4 mm (0.41 in) in mantle length and males 5.9 mm (0.23 in).[3] Average wet weights are around 0.20 and 0.02 g (0.00705 and 0.00071 oz), respectively.[3]

Other tiny species include members of thebobtail squid familySepiolidae; themyopsid squid generaAustraliteuthis andPickfordiateuthis; theoegopsid squid generaAbralia andAbraliopsis; the pygmy cuttlefishSepia pulchra; and the ram's horn squid,Spirula spirula.

Male dwarfism

[edit]

The octopod superfamilyArgonautoidea is characterised by markedly dwarfed males.[34] The four extant genera of the group areArgonauta,Haliphron,Ocythoe, andTremoctopus, all of which are exclusivelypelagic. The greatest disparity in the size of the sexes is seen in theblanket octopuses of the genusTremoctopus.Normanet al. (2002) reported a fully mature maleTremoctopus violaceus measuring 2.4 cm (0.94 in) in total length and weighing a mere 0.25 g (0.0088 oz). By comparison, the large females of this species reach total lengths of 2 m (6.6 ft) and probably some 10 kg (22 lb) in weight. This is the most extremesexual size dimorphism known among non-microscopic animals,[nb 3] with mature females being at least 10,000 times heavier than males, and likely up to 40,000 times heavier.[35] The related generaArgonauta andOcythoe have similarly small males, but the females are not nearly as large as those ofTremoctopus, and the size dimorphism is therefore less pronounced. The females of the fourth argonautoid genus,Haliphron, are the largest of all up to length of 3.5 m (11 ft) and mass of 75 kg (165 lb) (and possibly thelargest octopuses of any kind), but the males are also much larger, at up to 30 cm (12 in). So, the size dimorphism is least pronounced of all.[36]

Extinct taxa

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Numerous species of so-called micromorphic ammonites are known.[37]Maximites from theUpper Carboniferous is the smallest known ammonoid. Adult specimens reached only 10 mm (0.39 in) in shell diameter.[38]

Maximum size

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Scientifically validated records

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The largest cephalopod specimen ever recorded: a 495 kg (1,091 lb)colossal squid

Squids are the largest living cephalopods in terms of each ofmantle length, total length, and mass, with the largest species by at least two of these measures being thecolossal squid,Mesonychoteuthis hamiltoni. Reaching an estimated 3 m (9.8 ft) in mantle length and 10 m (33 ft) in total length, and weighing as much as 495 kg (1,091 lb), this species is also the largest of allextantinvertebrates.[39] The only other squid that approaches these dimensions is thegiant squid of the genusArchiteuthis, with females up to 275 kg (606 lb), 2.4 m (7.9 ft) in mantle length, and possibly as much as 15 m (49 ft) in total length, making it likely the longest of all cephalopods.[40] The two largestoctopus species—Enteroctopus dofleini andHaliphron atlanticus—can both exceed 70 kg (150 lb), and the former has a maximum total length of more than 6 m (20 ft).Cirrate (finned) octopods can also reach a large size, with the largest captured specimen likely being aCirrothauma magna of 1.7 m (5.6 ft) total length and 33 cm (13 in) mantle length,[41] though observations from submersibles suggest that members of this group can exceed 4 m (13 ft) in total length.[42]

Members of the other cephalopod groups are substantially smaller, although the largestcuttlefish can exceed 10 kg (22 lb) in weight and 50 cm (1.6 ft) in mantle length. Cephalopods of comparable size to the largest present day squid are known from fossil remains, including enormous examples ofammonoids,belemnoids,nautiloids,orthoceratoids,teuthids, andvampyromorphids.[43]

Colossal squid (Mesonychoteuthis hamiltoni)

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Beak of a colossal squid, which has the largest beak among living cephalopods

Though a substantial number ofcolossal squid (Mesonychoteuthis hamiltoni)remains have been recorded (Xavieret al., 1999 collated 188 geographical positions for whole or partial specimens caught by commercial and scientific fisheries), very few adult or subadult animals have ever been documented, making it difficult to estimate the maximum size of the species.McClainet al. (2015) stated that only 12 "complete" specimens were known.The largest known complete specimen of the colossal squid was a mature female captured in theRoss Sea in February 2007. Its weight was initially estimated at 450 kg (990 lb), its mantle length at 4 m (13 ft), and its total length at 8–10 m (26–33 ft).[44] Once completely thawed the specimen was found to weigh 495 kg (1,091 lb), but to measure only 2.5 m (8.2 ft) in mantle length and 4.2 m (14 ft) in total length.[45] It is likely that the specimen, and particularly its tentacles, shrank considerablypost mortem as a result of dehydration, having been kept in a freezer for 14 months. (As reported by theMuseum of New Zealand Te Papa Tongarewa, specimens ofNototodarus sloanii, the New Zealand arrow squid, can shrink by as much as 22% when dehydrated with alcohol solutions.[45]) The colossal squid specimen contracted by a further 5% after several years in preservative fluid (firstformalin and laterpropylene glycol).[46] The fins of the 2007 Ross Sea specimen measured around 1.2 m (3.9 ft) across and it had a mantle width of 98.2 cm (3.22 ft).[47] The arms ranged in length from 0.85 to 1.15 m (2.8 to 3.8 ft), while the two tentacles were around 2.1 m (6.9 ft) long.[48]

The 2007 Ross Sea specimen as it appears now, on display at theMuseum of New Zealand Te Papa Tongarewa

Beaks recovered from sperm whale stomachs indicate the existence of animals surpassing even the 2007 Ross Sea specimen. That specimen had alower rostral length (LRL) of 42.5 mm (1.67 in) and weighed 495 kg (1,091 lb),[49] whereas the 300 kg (660 lb) submature female from 2003 had a LRL of 37 mm (1.5 in).[50] By comparison, the largest known colossal squid beak from a sperm whale stomach measured 49 mm (1.9 in) in LRL.[22] Though the number of large colossal squid specimens known to science is too small to get a good idea of the relationship between beak size and overall body size, a beak of such great size indicates a truly massive animal weighing perhaps as much as 600–700 kg (1,300–1,500 lb).[49] However, thescaling relationship for this species shows considerable latitude, as demonstrated by a beak of 40 mm (1.6 in) LRL extracted from an animal weighing only 160 kg (350 lb).[49]

Giant squid (Architeuthis dux)

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A giant squid specimen; the extremely long feeding tentacles of the giant squid make it the longest known cephalopod, past or present.

The maximum size of thegiant squid (Architeuthis dux) has long been a subject of both popular debate and academic inquiry.[51] Unlike the colossal squid, the giant squid is known from a substantial number of mature specimens. Thetotal number of recorded specimens (across all developmental stages) approaches a thousand, with approximately 700 documented as of 2015[update], of which around 460 had been measured in some way.[52] This number has since increased substantially, with 57 specimens recorded from Japanese waters during an exceptional 15-month period between 2014 and 2015.[53]

A giant squid measuring 7 m (23 ft) in total length and weighing 250 kg (550 lb) displayed in a 3.5-tonne (7,700 lb) block of ice at theMelbourne Aquarium[54]

Based on a 40-year data set of more than 50 giant squid specimens,Roper & Shea (2013:114) suggest an average total length at maturity of 11 m (36 ft) and a "rarely encountered maximum length" of 14–15 m (46–49 ft). Of the nearly 100 specimens examined byClyde Roper, the largest was "46 feet (14 m) long".[55]O'Shea & Bolstad (2008) give a maximum total length of 13 m (43 ft) for females based on the examination of more than 130 specimens, measuredpost mortem and relaxed, as well as beaks recovered fromsperm whales (which do not exceed the size of those found in the largest complete specimens).Steve O'Shea estimated the maximum total length for males at 10 m (33 ft).[56] Older records of 18 m (59 ft) or more were likely exaggerated by stretching of the longfeeding tentacles or resulted from inadequate measurement methods such as pacing.[57] O'Shea has stated that, given the available evidence, the highestupper bound he would consider plausible for the giant squid's total length would be 15 m (49 ft), and that the likelihood that there exist 20-metre giant squid is "so exceedingly remote that you couldn't justify the effort in writing about it".[58]

Including the head andarms but excluding thetentacles (standard length, SL), the species very rarely exceeds 5 m (16 ft) according toO'Shea & Bolstad (2008).Paxton (2016a) considers 9.45 m (31.0 ft) to be the greatest reliably measured SL, based on a specimen reported byVerrill (1880a:192), and considers specimens of 10 m (33 ft) SL or more to be "very probable", but these conclusions have been criticised by giant squid experts.[59]O'Shea (2003a) put the maximum weight of female giant squid at 275 kg (606 lb), based on the examination of some 105 specimens as well as beaks recovered fromsperm whales (which do not exceed the size of those found in the largest complete specimens). Giant squid aresexually size dimorphic, with the maximum weight for males estimated at 150 kg (330 lb),[56] though heavier specimens have occasionally been reported (such as a 190 kg (420 lb) specimen[60] and a 163 kg (359 lb) specimen[61]).Roper & Jereb (2010a:121) give a maximum weight of up to 500 kg (1,100 lb), and "possibly greater". Discredited weights of as much as atonne (2,200 lb) or more are not uncommon in older literature (seebelow).[22]

Other squid taxa

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Dana octopus squid

The third-heaviest extant squid species isTaningia danae, also known as the Dana octopus squid. The largest well documented specimen is a 160 cm ML mature female reported byRoper & Vecchione (1993) from the North Atlantic. The original paper consistently gave the mass of this specimen as 61.4 kg (135 lb), but according toRoper & Jereb (2010h:266) this figure is wrong and stems from a typographical error, the correct mass being 161.4 kg (356 lb). Another similarly large specimen—a female weighing 124 kg (273 lb)—was reported from northern Spanish waters byGonzálezet al. (2003:297) (see also initial reports[62]). In July 2010, asperm whale was photographed off theAzorean island ofFaial with a large squid—likelyT. danae—in its mouth. The specimen's maximum width, from fin tip to fin tip, was estimated at 1.5–2 m (4.9–6.6 ft); this would approximate its mantle length.[63]

Onykia robusta grows to a mantle length of at least 2 m (6.6 ft)

Onykia robusta, previously known asMoroteuthis robusta[64] and sometimes called the robust clubhook squid, has a mantle length of up to 200 cm (6.6 ft).[65] Some older records exceed this, such as the 91.5 in (232 cm) ML reported byVerrill (1876:237) from a specimen with a total length of 14 ft (4.3 m) (excluding the ends of the tentacles, which had been destroyed).Nesis (1987:192) likewise gave a maximum mantle length of 230 cm (7.5 ft), butRoper & Jereb (2010i:364) wrote that "this old record might be in error", with the species commonly growing to 160 cm (5.2 ft) ML.Glaubrecht & Salcedo-Vargas (2004:66) provided a maximum total length of 4–6 m (13–20 ft). Literature sources give a maximum weight of 50 kg (110 lb).[66] There exist numerous published records of large individuals of this species.[67]

Fisherman taking aHumboldt squid off the coast ofChile. This species is the largest squid species commonly encountered by humans, possessing a large fishery.[68]

TheHumboldt squid (Dosidicus gigas), also known as the 'jumbo squid', grows to a maximum mantle length of at least 120 cm (3.9 ft),[69] if not 150 cm (4.9 ft).[70] The largest animals are found off the western coast ofSouth America; northern populations reach 100 cm (3.3 ft) ML, and in general 50–80 cm (1.6–2.6 ft) ML is more typical for the species.[71] Southern populations may have a total length approaching 2.5 m (8.2 ft),[71] and possibly up to 3.7 m (12 ft).[72] Again, specimens from theNorthern Hemisphere are much smaller, with those off theCalifornian coast reaching total lengths of less than 1.7 m (5.6 ft).[73] The Humboldt squid commonly attains a weight of around 20–30 kg (44–66 lb)[71] and can reach a maximum of 50 kg (110 lb).[74] There are anecdotal reports of much larger individual animals, including from diverScott Cassell, who has dived with Humboldt squid over 300 times (reportedly more than any other person).[75][nb 4]

Moroteuthopsis longimana, previouslyKondakovia longimana and sometimes known as the giant warty squid, is a little known species with a circum-Antarctic distribution in theSouthern Ocean. The largest complete specimen, found floating at the surface off theSouth Orkney Islands, had a mantle length of 108 cm (3.54 ft),[76] but a damaged female specimen with an estimated mantle length of around 150 cm (4.9 ft) is known.[77] The largest complete specimen had a wet weight of 29 kg (64 lb).[76] The species' maximum weight has been estimated at 50 kg (110 lb).[78]

Largest octopodes

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The giant Pacific octopus is one of the two largest octopus species

Thegiant Pacific octopus (Enteroctopus dofleini) grows to more than 6.1 m (20 ft) in total length[79] and at least 60 cm (2.0 ft) in mantle length.[80]Cosgrove (1987) andCosgrove & McDaniel (2009:69) gave a maximum confirmed weight of 71 kg (157 lb) for a live specimen collected in the mid-1960s.[40]Normanet al. (2014:124) accept a maximum weight of at least 180 kg (400 lb), which approximates the 182.3 kg (402 lb) reported for a specimen caught offSanta Barbara,California, in 1945, of which photographic evidence survives.[81] No specimens approaching this size have been reported since the middle of the 20th century, with recent specimens very rarely exceeding 50 kg (110 lb).[82] It is possible that the maximum size of the species has decreased over this period, perhaps due tobioaccumulation oftoxicants (seebelow).[83]

In 2002, a giant specimen ofHaliphron atlanticus, the seven-arm octopus, was caught by fishermen trawling at a depth of 920 m (3,020 ft) off the easternChatham Rise,New Zealand. This specimen, the largest of this species and possibly of all octopuses, was the first validated record ofHaliphron from theSouth Pacific. It had a mantle length of 0.69 m (2.3 ft), a total length of 2.90 m (9.5 ft), and a weight of 61.0 kg (134.5 lb), although it was incomplete.[84] The total length of the specimen, when complete, has been estimated at 4 m (13 ft), and its weight at 75 kg (165 lb).[85]

Extinct taxa

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The largest known specimen ofParapuzosia seppenradensis, with its describing author,Hermann Landois, who reconstructed its missing living chamber with a combination of wire frame and paper.[86] Landois assumed that the living chamber constituted one-fourth of the outerwhorl, butTeichert & Kummel (1960:6) estimated that it continued for three-fourths to one full whorl, which would give an original shell diameter of around 3.5 m (11 ft).

Certain extinct cephalopods rivalled or even exceeded the size of the largest living species.[87] In particular, the subclassAmmonoidea is known to have included a considerable number of species that may be considered "giant" (defined byStevens, 1988 as those exceeding 1 m (3.3 ft) in shell diameter). The largest confirmedammonite, a specimen ofParapuzosia seppenradensis discovered in aGerman quarry in 1895, measures 1.742 m (5.72 ft) in diameter,[88] though itsliving chamber is largely missing. The diameter of the complete shell has been estimated at 2.55 m (8.4 ft), assuming the living chamber took up one-fourth of the outerwhorl.[89]Teichert & Kummel (1960:6) suggested an even larger original shell diameter of around 3.5 m (11 ft) for this specimen, assuming the body chamber extended for three-fourths to one full whorl. In 1971 a portion of an ammonite possibly surpassing this specimen was reportedly found in abrickyard inBottrop, western Germany.[90] A specimen found by Jim Rockwood, from theLate Triassic nearWilliston Lake,British Columbia, was said to measure more than 8 ft (2.4 m) across, but was later determined to be aconcretion.[91]

Reconstructions of theholotype (FHSM 13049) and largest specimen (NDGS 241) ofEnchoteuthis melanae, based onMuensterella andDosidicus

Heteromorph ammonites are known to have exceeded 1 m (3.3 ft) in length also, but since their shells were uncoiled to varying degrees, they were overall much smaller than the largest non-heteromorphs. The greatest lengths of all were reached by theorthocones ofendoceridnautiloids such asEndoceras, which may have exceeded 8 m (26 ft), although their maximum size is uncertain; while the largest well documented endocerid fossil is likely the 3-metre-long (9.8 ft) shell fragment housed at theMuseum of Comparative Zoology,Harvard University, there are published reports of even larger specimens.Teichert (1927) mentioned specimens up to 5 m (16 ft) long from theMiddle Ordovicianlimestone ofEstonia[92] andFrey (1995:72) gave a maximum shell length of 6 m (20 ft) for the group. On the subject of endocerid size, nautiloid specialistRousseau H. Flower wrote:

They are not all large, by any means, but specimens twelve feet [3.7 m] in length have been collected, and fragments of greater diameter indicate a much greater maximum length. I am not wholly inclined to discredit a report of an endoceroid found in a quarry nearWatertown New York, which was measured before it was broken up and found to attain a length of 30 feet [9.1 m].[93]

However, the uncoiled length of the largest ammonites far exceeds that of even these giant endocerids.Parapuzosia seppenradensis, the largest known ammonite species, had an estimated maximum unrolled shell length of around 18 m (60 ft). It was also possibly the heaviest of all known cephalopods, past or present, with an estimated live mass of 1,456 kg (3,210 lb), of which the shell would constitute 705 kg.[94] By comparison, the largest endocerids may have weighed around 1,000 kg (2,200 lb).[7] In terms of mass, these are the largest knowninvertebrates that have ever lived,[95] though perhaps still second to the largest living cephalopods when consideringtissue mass alone, since in shell-bearing species the vast majority of the living tissue is restricted to thebody chamber, which occupies only a fraction of the internal shell volume.[8] They might also be the largest—or at least longest—shell-bearing animals that have ever lived.[8][nb 5]

CretaceousYezo Group inJapan yields multiple taxa of large-sized cephalopod fossil remains.OegopsidYezoteuthis giganteus andHaboroteuthis poseidon are only known from jaw elements, which is close to size of that of giant squid. Species ofNanaimoteuthis, especiallyN. hikidai would be the largest known vampyromorph.[96] Unnamed, incomplete upper jaw fossil that is approximately twice as large as that of the mature giant squid is also found there.[97]

Historical claims

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Misidentifications

[edit]
Kubota's photograph of a largeOnykia robusta, which was misidentified as thegiant squid,Architeuthis dux.Wide-angle photography exaggerated the size of the squid[98] such that it appeared impossibly large for the species.

The maximum sizes of certain cephalopod species, most notably the giant squid and giant Pacific octopus, have often been misreported and exaggerated. The literature on cephalopod size has been further muddied by the frequent misattribution of various squid specimens to the giant squid genusArchiteuthis, often based solely on their large size. In the academic literature alone, such misidentifications encompass at least theoegopsid familiesChiroteuthidae,Cranchiidae,Ommastrephidae,Onychoteuthidae, andPsychroteuthidae.[99][nb 6] This situation is further confused by the occasional usage of thecommon name 'giant squid' in reference to large squids of other genera.[100]

Perhaps the most notable misidentification relates to a photograph taken some time before 1993 by diver H. Kubota off southernJapan.[101] The image shows a large individual ofOnykia robusta (previously known asMoroteuthis robusta), which appears to be sick or dying, alongside a diver in shallow water.[98] A video of the same animal appeared in a Japanese made-for-television film.[98] The image was published in the 1993 bookEuropean Seashells byGuido T. Poppe and Yoshihiro Goto, where it was identified asArchiteuthis dux, the giant squid, and said to have been taken in theNorth Atlantic.[102] If true, this image would represent the first known photograph of a live giant squid. InThe Search for the Giant Squid (1998),Richard Ellis wrote:

For a moment, I thought that some obscure photograph had captured the most elusive image in natural history. Fortunately for those who have devoted their lives to searching forArchiteuthis, this was only an aberration, a case of mistaken identity.[98]

It would be more than a decade before the true first photographs of a live giant squid in the wild were taken on 30 September 2004 byTsunemi Kubodera andKyoichi Mori.[103] Kubodera and his team subsequently became the first tofilm a liveadult giant squid on 4 December 2006,[104] and the first tofilm a live giant squidin its natural habitat in July 2012.[105] These milestones were preceded by the first footage of a live (paralarval) giant squid in 2001,[106] and the first image of a live adult giant squid on 15 January 2002.[107] Since then, live giant squid have been photographed and filmed on a number of occasions.[108]

Giant squid

[edit]
The "nearly perfect specimen" that was beached alive inTrinity Bay,Newfoundland, on 24 September 1877. A number of exceptionally large giant squid were reported from Newfoundland in the 1870s, and these were meticulously documented in a series of papers by zoologistAddison Emery Verrill.[109]

Reports of giant squid (Architeuthis dux) specimens reaching or even exceeding 18 m (59 ft) in total length are widespread, but no animals approaching this size have been scientifically documented in recent times.[110] This is despite there beinghundreds of specimens available for study (c. 700 documented as of 2015, of whichc. 460 measured in some way[52]), including numerous recent examples, such as the 57 specimens recorded from Japanese waters over a 15-month period in 2014–2015.[53] It is now thought likely that such lengths were achieved by great lengthening of the two longfeeding tentacles, analogous to stretching elastic bands, or resulted from inadequate measurement methods such as pacing.[57]

On the subject of the oft-cited maximum size of 18 metres—or 60 feet—Dery (2013) quoted giant squid expertsSteve O'Shea andClyde Roper:

If this figure [45 ft or 14 m] seems a little short of theBrobdingnagian claims made forArchiteuthis in mostpop-science stories about the animal, that's probably because virtually every general-interest article dutifully repeats the magic number of60 feet.

Steve O'Shea deplores the media's perpetuation of what he believes to be a credulity-straining exaggeration, based on the 19th-century biologistThomas Kirk's eyeball estimate of a specimen's length.[nb 7] In a comment on the final draft of this article, O'Shea wrote, "Kirkpaced it, in his own words, for he had no ruler/measure handy, and I believe this misrepresentation has been perpetuated enough; if they were foot-on-foot, as in heel directly to toe, I would accept 57 (or 58, whatever the precise figure was), but I think perpetuating this as fact any longer is doing a disservice to science."

Roper, in his comments on the final draft of this article, was even more conservative, writing, "there are noconfirmed records of giant squid longer than about 45 feet [14 m] total length. Most are in the 25–35-foot [7.6–10.7 m] range. I have examined specimens in museums and laboratories around the world—perhaps a 100 or so—and I believe the 60-foot number comes from fear, fantasy, and pulling the highly elastic tentacles out to the near breaking point when they are measured on the shore or on deck."

Largest reported animals
[edit]
Frequency distribution of total length, mantle length, and mass inArchiteuthis dux. The 2,000 lb (910 kg) extremeoutlier is sourced fromVerrill (1880a) and is unlikely to be accurate; the next most massive individual in the data set was only 700 lb (320 kg), and 95% of specimens were below 250 kg (550 lb).[40]
Log–loglinear regressions forArchiteuthis dux: (A) total length vs. mass; (B) mantle length vs. total length; and (C) mantle length vs. mass[40]

Paxton (2016a) investigated the maximum size ofArchiteuthis by performing a statistical analysis using data fromliterature records of giant squid specimens. He selected what he regarded as the largest size records for each of mantle length (ML), standard length (SL), and total length (TL). Paxton's study has been criticised by giant squid experts, who have called into question the reliability of some of the selected literature records.[59]

For mantle length,Paxton (2016a:83) considered the 11 ft (3.35 m) reported byDell (1952:98) as the "longest measured", though "more reliably" the 9 ft 2 in (2.79 m) ML specimen fromLyall Bay,New Zealand, documented byKirk (1880:312).[nb 8] Paxton added: "A 4.5 m [15 ft] specimen fromMauritius is often mistakenly cited but consultation of the primary paper (Staub, 1993) reveals an ill-defined length which is clearly not ML." The greatest measured ML of a giant squid recovered from asperm whale is either the 2.4 m (7.9 ft) reported byKeil (1963:320) (though Paxton writes: "the account is confused and the 2.4 m figure probably refers to the head and ML combined") or the 6 ft 6 in (1.98 m) of a specimen that had been swallowed whole off theAzores, detailed byClarke (1955:589) andClarke (1956:257). The "longest visually estimated" ML, according to Paxton, is thec. 100 ft (30 m) of a specimen apparently observed in theNorth Atlantic offPortugal, attributed to a personal communication with T. Lipington. A more modest 4 m (13 ft) ML is also given, based on a sighting in theIndian Ocean sourced to the TV documentary ofLynch (2013).

For standard length (excluding thetentacles),Paxton (2016a:83) cited the 31 ft (9.45 m) of the "Three Arms specimen" documented byVerrill (1880a:192) as the "longest measured". Among specimens recovered from sperm whales, the longest "definitely measured" SL is the 16 ft 3 in (4.95 m) reported byClarke (1956:257) and the longest "visually estimated" SL is thec. 9 m (30 ft) attributed to a photograph of a sperm whale with giant squid remains in its jaws,[111] though Paxton conceded that it is "[n]ot clear how much/what portion of body was eaten". For the "longest visually estimated", more extreme supposed SLs ofc. 175 ft (53 m) andc. 100 ft (30 m) are cited toStarkey (1963) andEllis (1998a:246), respectively (the latter an eyewitness account by Dennis Braun). Paxton treated these last two size estimates as SLs as opposed to TLs because "squid do not generally leave their tentacles exposed except when grabbing prey and this appears to be the case forArchiteuthis".

For total length,Paxton (2016a:83) considered three records as candidates for the "longest measured": the 19 m (62 ft) specimen ofBerzin (1972:199), the 55 ft 2 in (16.81 m) specimen described byKirk (1888) asArchiteuthis longimanus—a strangely proportioned animal that has been much commented on[nb 7]—and the 55 ft (16.76 m) "Thimble Tickle specimen" reported byVerrill (1880a:191), which is often cited as the largest giant squid ever recorded.[nb 9] Of the last one, Paxton wrote: "Sometimes mistakenly cited as 17.37 m (57.0 ft) but the source is clear that it is 55 ft long." The first two records, particularly that of Berzin, are more questionable, as Paxton explained:

The accuracy of the two longest measured TLs of 19 and 16.81 m from a specimen found in the gut of a sperm whale from the Indian Ocean and from the specimen from New Zealand in 1887, respectively, should also be questioned but again are certainly not impossible. The New Zealand specimen (namedArchiteuthis longimanusKirk, 1888) clearly has the largest ratio of TL to ML ever known inArchiteuthis [...] which led [O'Shea & Bolstad, 2008] to suggest that the length was paced out and/or there was extensive post-mortem stretching. However, a re-reading of the original paper suggests that the specimen, although initially paced out, was actually measured, nevertheless the TL is at the edge of the 99.9% prediction interval range [...] and so it was certainly an unusual specimen.Berzin's (1972) Indian Ocean claim is suspect because of the roundness of the figure, the lack of detailed measurements and because in an associated photo, the mantle (whose length was not given) does not look very large compared to the men in the image. Consequently the measurement, if accurate, would represent another animal with very long tentacles.[117]

The 19-foot (5.8 m) tentacle that fisherman Theophilus Picot hacked off a live animal on 26 October 1873. Picot estimated the total length of the squid at 60 ft (18 m).[118]

However, asPaxton (2016a:86) pointed out, the genetic analysis ofWinkelmannet al. (2013)—which concluded that there is likely a single, globally-distributed species ofArchiteuthis—did not encompass these two specimens, and it is therefore possible that there exists a second, as yet unsampled, giant squid species with proportionately longer tentacles.

The 19 m (62 ft) total length of the Berzin specimen was later confirmed to be erroneous; according to Valentin Yukhov, who was involved in the specimen's discovery, it should have read 9 m (30 ft).[119] The misprint was reproduced in the English translation published the following year and was later propagated in a number of papers on giant squid.[119] With the Berzin specimen not being as large as originally reported, the longest giant squid recovered from a sperm whale is the 34 ft 5 in (10.49 m) TL individual recorded byClarke (1956:257) (this specimen also has the longest confirmed ML and SL of any giant squid from a sperm whale).[120] Paxton considered the "longest visually estimated" TL to be the 60 ft (18 m) published byMurray (1874:121), from an eyewitness account by fisherman Theophilus Picot, who claimed to have struck the floating animal from his boat,causing it to attack. Picot managed to hack off one of its tentacles, which was subsequently examined by a number of authors.[121]

Perhaps the largest of all recorded giant squid specimens was the one found floating at the surface offSaint-Gilles,Réunion, on 4 March 2016.[119] Although due to its great size the specimen could not be retrieved in its entirety, the head and arm crown were saved. Crucially, this meant the beak could be measured to estimate the mantle length and total length of the specimen. Using differentallometric scaling equations,[nb 10] thelower rostral length of the beak, at 19.74 mm (0.777 in), gave an estimated dorsal mantle length of 215.3–306.0 cm (7.064–10.039 ft) and this, in turn, was used to estimate the total length at 11.025–15.664 m (36.17–51.39 ft).[119]

Supposed sucker scars
[edit]
A dramatisation of an underwater encounter between the sperm whale and giant squid, from adiorama at theAmerican Museum of Natural History

More extreme and outlandish giant squid size claims—belonging firmly in the realm ofcryptozoology—have appeared in the works of authors such asBernard Heuvelmans,Willy Ley, andIvan T. Sanderson.[126] The existence of these gargantuan squids is often supported by reference to the giant circular scars sometimes found onsperm whales, which are assumed to have been inflicted by thesuckers of struggling giant squid. Sometimes these claims are accompanied by extrapolations of body size based on theisometric upscaling of a "typical" giant squid.[127] However, such scars are not necessarily of squid origin and may instead represent fungal growths or bite marks, withsea lampreys (Petromyzon marinus) being one possible source.[128] Even in the case of genuine giant squid sucker marks it is possible that subsequent skin growth has enlarged them well beyond their original dimensions.[129]

A portion ofsperm whale skin with giant squid sucker scars. In this widely reproduced image the largest sucker scars measure around an inch (2.5 cm) across.[130]

Nevertheless, claims of enormous sucker scars are widespread in the literature.Richard Ellis collected some of "the more egregious examples" in his book,The Search for the Giant Squid.[131] These include the claim ofDozier (1976) that "an ordinary giant squid of 50 feet [15 m] leaves teeth-ringed sucker marks measuring between three and four inches [7.6–10.2 cm] across on a whale, but sperm whales have been captured with tentacle marks 18 inches [46 cm] across."L. Harrison Matthews's monographic treatment of the sperm whale, published in 1938, includes the following: "Nearly all male Sperm whales carry scars caused by the suckers and claws of large squids, scars caused by suckers up to 10 cm. [3.9 in] in diameter being common. The claw marks take the form of scratches 2–3 m. [6.6–9.8 ft] in length, and appear to be of more frequent occurrence than sucker marks".[132]Ellis (1998a:142) wrote that this 10 cm figure is "so much larger than any other recorded sucker dimensions that one suspects some sort of error, either in measuring or in transcription."

The subject was covered in some detail byWood (1982:192):

Measurements of 90 ft [27 m], 130 ft [40 m] and even 200 ft [61 m] have been conjectured for giant squids from the size of sucker marks found on the skins of captured sperm whales, but it is dangerous to place too much reliance on this evidence.Verrill says the largest suckers on the tentacles of a 32 ft [9.8 m] long specimen measured 114 in [3.2 cm] in diameter, and those on a 52-footer [16 m] about 2 in [5.1 cm].Daniel (1925), however, examined sucker marks on the head of one cachalot which measured 312 in [8.9 cm] across, and others measuring up to 5 in [13 cm] in diameter have been found on the skins of sperm whales captured in the North Atlantic.Ivan Sanderson (1956) goes even further and claims that sucker marks over 18 in [46 cm] have been found on the heads of cachalots, but he does not explain how the poor whales managed to escape from the clutches of such colossi!

The general consensus of opinion is that exceptionally large sucker marks, i.e. over 2 in [5.1 cm] in diameter, are old scars that have increased in size as the sperm whale grew.

One of the pairedtentacular clubs ofArchiteuthis, showing the enlarged suckers of the manus, which are the largest found on any of the giant squid'slimbs

Perhaps the most extreme published claim, ridiculed byEllis (1998a:142), appeared inWilly Ley's 1959 book,Exotic Zoology: "Toothed whales, vomiting in death struggle, have shown evidence of still largerkraken; in one case a 6-foot [1.8 m] piece of tentacle,with a diameter of 2 feet [0.6 m; emphasis in original], has been claimed. Another claim goes for marks on the skin of such a whale, looking like the mark of a sucking disk over 2 feet [0.6 m] in diameter".[133]

Marine biologistFrederick Aldrich, who personally examined more than a dozen giant squid specimens, wrote that his largest specimen fromNewfoundland bore tentacular suckers "approximately two inches [5.1 cm] in diameter" but that "[s]uckers and their toothed armament of over twelve inches [30 cm] in diameter have been found in the stomachs of sperm whale as indigestible wastes".[134] This led him to entertain the idea of giant squid over 150 ft (46 m) long,[nb 11] and even to suggest abinomial name for this super-sized species, were it ever to be discovered:Architeuthis halpertius.[134]

By comparison, giant squid suckersnormally reach a maximum diameter of only a few centimetres. Based on a detailed examination of a number of large specimens from New Zealand waters,Förch (1998:55) wrote that "[t]he largest suckers [...] on the sessile arms are a very constant 21–24 millimetres [0.83–0.94 in] in external diameter". In giant squid the largest suckers of all are found on the central portion of thetentacular club, called themanus, and among the specimens examined byFörch (1998:53) these reached a maximum diameter of 28–32 mm (1.10–1.26 in).Clarke (1980) wrote: "I have not yet seen conclusive evidence to suggest that sucker scars are larger than 3.7 centimetres [1.46 in] across".[138] According toRoper & Boss (1982:97), the largest suckers of the tentacular clubs reach 5.2 cm (2.0 in) in diameter.

Mass estimates
[edit]
A dozen people attempting to move agiant squid weighing in excess of 200 kg (440 lb)[139]

It is now accepted that the giant squid hasa maximum mass of several hundred kilograms,[140] but the literature is rife with claims of much greater weights.Clarke (1966), for example, put the mass of the largest giant squid specimens at around 1 tonne (2,200 lb).[141] Similarly,Richard Ellis wrote: "Where [giant] squid carcasses have actually been weighed, it appears that the longest ones—in the 50-foot [15 m] range, for example—weigh about a ton [910 kg]."[142] Much greater estimates of giant squid mass can be found in, for example,Natural History of Marine Animals byMacGinitie & MacGinitie (1949): "two arms ofArchiteuthis that were 42 feet [13 m] long were found, and if one reconstructed a body [...] the squid to which these arms belonged was 4.6 feet [1.4 m] in diameter and 24 feet [7.3 m] long, with an overall measurement of 66 feet [20 m]. It would have weighed about 4212 tons [38.6 tonnes]." They added that a 55-foot (17 m) specimen, such as the one reported fromThimble Tickle,[nb 9] "would have weighed 2914 or 30 tons [26.5 or 27.2 tonnes] including the tentacles—a truly noble animal, being a little more than one-fifth the weight of thelargest whale and larger than thewhale sharks andbasking sharks, the largest of all fishes".[143] Ellis characterised these estimates as "unfounded exaggerations".[142] In the revised edition ofNatural History of Marine Animals, published in 1968, the authors reduced their estimate to less than 8 tonnes.[144]

Bernard Heuvelmans believed that "there must beArchiteuthis weighing more than 5 tons, and some even larger ones which must weigh between 2 and 27 tons, the normal weight being around 8 tons. There are good reasons to believe that there may even exist specimens twice as long as that of Thimble Tickle, which, depending upon their girth, might have weighed between 16 and 216 tons, but more likely around 64 tons."[145] Ellis, who considered these estimates "utterly ridiculous", wrote:

Heuvelmans commits a fundamental error in calculating the weight of some of these monsters when he writes that "the density of living creatures is only slightly higher than that of water ... a decimetre of living flesh weighs about as much as a litre of water." That may be true for some other living creatures, but the flesh ofArchiteuthis, saturated withammonium chloride, islighter than water, and the giant squid isneutrally buoyant. (This is believed to be the reason that dead or dying squid are found floating at the surface or washed up on the beach.) His assumption, therefore, that the 55-foot-long [17 m] Thimble Tickle squid would have "probably weighed near 24 tons" is patently erroneous.[146]

On the subject of the Thimble Tickle specimen's mass,Wood (1982:190) referred to the work ofSoviet zoologist and writerIgor Akimushkin:

According to DrIgor Akimushkin (1965), the Russian teuthologist, a 12 m [39 ft] long giant squid will weigh 1 tonne [2,200 lb] if the head, mantle and arms combined make up half the total length. Since there is acubic relationship between the linear dimensions ofArchiteuthis and its volume or weight, this means the Thimble Tickle monster must have scaled about 2.8 tonnes [6,200 lb] (i.e. the weight of a large bullhippopotamus), although 2 tonnes [4,400 lb] is probably a more realistic figure.

Giant Pacific octopus

[edit]

The maximum size of thegiant Pacific octopus (Enteroctopus dofleini) has long been a source of debate in the scientific community, with dubious reports of specimens weighing hundreds of kilograms.

Largest reported animals
[edit]
Frequency distribution of mass and interocular distance in male and femaleEnteroctopus dofleini[40]
Log–log plot showing relationship between interocular distance and mass inEnteroctopus dofleini[40]

In 1885, reporting on the longest octopus specimen reliably recorded up to that point, renownedmalacologistWilliam Healey Dall wrote:

In 1874 I speared an octopus in the harbor ofIliuliuk,Unalashka, which was afterward hung, by a cord tied around the body immediately behind the arms, to one of thesterndavits of the coast survey vessel under my command. As soon as the animal died and the muscles relaxed, I noticed that the tips of the longer tentacles just touched the water. On measuring the distance with a cord, I found it to be sixteen feet [4.9 m], giving the creature a spread from tip to tip of the longest pair of arms, of not less than thirty-two feet [9.8 m]. The arms toward the tips were all exceedingly slender, but rather stout toward the body, which was somewhat over a foot [30 cm] long. The largest suckers were two and a half inches [6.4 cm] in diameter; the whole creature nearly filled a large washtub. Parts of this specimen are now in theU. S. national museum.[147]

In an article for theNational Marine Fisheries Service summarising knowledge on the giant Pacific octopus,High (1976:17–18) wrote:

Several octopuses in excess of 100 pounds [45 kg] have been encountered and captured. Much larger ones have been reported but, like theLoch Ness Monster, these usually elude the careful photographer or scientist. Most octopuses weigh less than 70 pounds [32 kg] with a stretched length of 15 feet [4.6 m] or less. Overall length between arms is not a suitable measure because of the animal's unusual elasticity.

In the late 1950s, I interviewed a Canadian commercial diver, Jock MacLean ofPrince Rupert,B.C. He reported capturing an immense creature weighing 600 pounds [272 kg] and measuring 32 feet [9.8 m] from arm tip to top. MacLean's photographs, unfortunately, were of poor quality. Smaller animals, to 400 pounds [181 kg], were occasionally taken in his commercial octopus fishing endeavor.

Hochberg & Fields (1980:436) referenced the same specimen, writing: "the largest specimen on record with a total arm spread of 9.6 m [31 ft] and a weight of 272 kg [600 lb]". These figures are only estimates, however, as—contrary to the above quotation fromHigh (1976:17–18)—it appears that this specimen was never collected and measured.[40] Murray Newman, director of theVancouver Aquarium for 37 years, quoted Jock MacLean in his 1994 memoir,Life in a Fishbowl: "Next year [1957] in the same place, I saw one, maybe thirty-two feet [9.8 m] across and six hundred pounds [272 kg]. Didn't go for her, though, no place to keep her!"[148] Nevertheless, themisleadingly precisemetricconversion of 272 kg (for 600 lb) and theimprecise conversion of 9.6 m (for 32 ft; naively employing aconversion factor of 0.3 instead of0.3048) gained wide acceptance as the maximum recorded dimensions of the giant Pacific octopus, and have been much repeated.[nb 12]

Jock MacLean is also reported to havecaptured a 198 kg (437 lb) animal with an arm span of 8.5 m (28 ft) nearPort Hardy,British Columbia, in March 1956.[150] Another giant specimen was caught offSanta Barbara,California, in 1945. Its weight was recorded as 182.3 kg (402 lb) and the surviving photograph makes it possible to estimate its total length at more than 3 m (9.8 ft) and arm span at 6–6.7 m (20–22 ft).[81] In a book dedicated to the giant Pacific octopus,Cosgrove & McDaniel (2009:72) summarised knowledge on the species's maximum size as follows:

The specimen William Dall speared in 1885 [sic] at Iliuliuk had the largest radial span of any giant Pacific octopus ever measured. Jock MacLean's 1956 Port Hardy behemoth was the biggest ever weighed. The Santa Barbara specimen photographed in 1945 was the second heaviest. It would appear that octopuses weighing as much as 272 kg (600 lb) and with radial spans of over nine metres (30 feet) are within the realm of possibility, but have never actually been documented by both measuring and weighing.

Possible diminution in size
[edit]

No specimens approaching these extreme sizes have been reported since the middle of the 20th century. This lack of giant individuals is corroborated by commercial octopus fishers; none of those interviewed byCosgrove & McDaniel (2009) had caught a single animal weighing more than 57 kg (126 lb) in the previous 20 years, among many thousands harvested over that period.[82] Octopus specialist Roland Anderson, a biologist with theSeattle Aquarium for more than 30 years, had long sought, unsuccessfully, to find a giant Pacific octopus weighing more than 100 lb (45 kg). In an attempt to raise a truly enormous specimen, Anderson fed a number of captive malesad libitum. The heaviest animal (nicknamed 'Big') attained a peak weight of 43 kg (95 lb) and its largest suckers measured 7.9 cm (3.1 in) in diameter.[151] Anderson suggested the species might now be maturing at a smaller size as a result oftoxicantbioaccumulation, which could explain the lack of truly gigantic specimens in recent times. In particular, high concentrations ofheavy metals andPCBs have been identified in thedigestive glands of wild giant Pacific octopuses, likely originating from their preferred prey, thered rock crab (Cancer productus).[152] A preliminary study found that aquarium animals fed equal quantities of raw sea food and liveC. productus (caught locally inElliott Bay) matured at a smaller size, reached a lower maximum weight (27 kg [60 lb] mean), and had higher concentrations of most heavy metals, than those fed solely on raw sea food (36 kg [79 lb] mean, including the aforementioned 43 kg [95 lb] specimen).[153]

Largest species by measure

[edit]

Cephalopod size can be quantified in various ways. Some of the most common size measures are covered below. The following four tables list only extant species;extinct taxa are treated separately at the end.

Mantle length

[edit]
TeuthologistClyde Roper lying alongside a largegiant squid (Architeuthis dux) specimen of almost 2 m (6.6 ft) ML
Reaching a mantle length of 2 m (6.6 ft),Onykia robusta is one of the largest squid species (the specimen shown here is considerably smaller).
A 52 lb (24 kg)Humboldt squid (Dosidicus gigas) caught off the southernCalifornian coast displaying deep redchromatophoric colouring.
Enteroctopus dofleini at theNational Aquarium inWashington, D.C.
Sepia apama, the largest species of cuttlefish, is native to the southern coast of Australia.
Sepia pharaonis, a large and commercially important cuttlefish species from theIndian Ocean
Bobtail squids, such as thisEuprymna berryi fromEast Timor, are some of the smallest of all cephalopods and are not known to attain a mantle length in excess of 10 cm (3.9 in).

The list of largest cephalopods by mantle length is dominated by squids, with more than twenty species exceeding the largest-bodied octopuses and cuttlefish. The largest of all is thecolossal squid (Mesonychoteuthis hamiltoni) with an estimated maximum mantle length of 3 m (9.8 ft) (Roper & Jereb, 2010c:173). Even greater mantle lengths have historically been reported for thegiant squid (Architeuthis dux), but these have been discredited (seeO'Shea & Bolstad, 2008).

Oegopsida (oegopsid squids)
SpeciesMaximum mantle lengthReferencesNotes
Mesonychoteuthis hamiltoni
(colossal squid)
≈300 cm (estimate)Roper & Jereb (2010c:173)The largest complete specimen, a mature female recovered from theRoss Sea in February 2007, had a mantle length of around 2.5 m ([Anonymous],N.d.)—this being the maximum scientifically documented mantle length (Rosaet al., 2017)—andseveral other specimens near this size have been recorded. However, at 42.5 mmLRL, itsbeak is considerably smaller than the largest recovered from asperm whale stomach (49 mm LRL;O'Shea & Bolstad, 2008;[Anonymous],N.d.). Maximum mantle lengths as great as 4 m have been estimated and reported in the past (Young, 2003;O'Shea & Bolstad, 2008). There are published claims of a very large section ofgladius that would suggest a colossal squid measuring 5 m or more in mantle length (Wood, 1982:191;Bright, 1989:146).[nb 13]
Galiteuthis phyllura? 265–275 cm (estimate)Nesis (1985);Nesis (1987:274);Ellis (1998a:149);Glaubrecht & Salcedo-Vargas (2004:65);Hoving & Robison (2017:47)Estimate based on 40 cm longarm and 115 cmtentacle from theSea of Okhotsk.[nb 14]Roper & Jereb (2010c:165) write: "this is considered a doubtful record that might refer to total length; probably the maximum mantle length is less than 400 to 500 mm".
Architeuthis dux
(giant squid)
240 cm (female)Landmanet al. (2004:686);O'Shea (2006);Roper & Shea (2013:114)Dorsal mantle length of female captured offTasmania,Australia, reported byLandmanet al. (2004:686) and cited byRoper & Shea (2013:114). Questionable records of up to 500 cm ML can be found in older literature (Roper & Jereb, 2010a:121).O'Shea & Bolstad (2008) give a maximum mantle length of 225 cm based on the examination of more than 130 specimens, as well as beaks recovered fromsperm whales (which do not exceed the size of those found in the largest complete specimens).Paxton (2016a) accepts a maximum recorded ML of 279 cm, based on theLyall Bay specimen reported byKirk (1880:312), but this record has been called into question as thegladius of this specimen—which should approximate the mantle in length—was said to be only 190 cm long (Greshko, 2016).[nb 8]
Onykia robusta
(robust clubhook squid)
200 cmNorman (2000:174);Bolstad (2008:107);Okutani (2015b)Kuboderaet al. (1998) give a maximum of at least 161.5 cm ML. The largest specimen seen byBolstad (2008:107) had a mantle length of 197 cm (USNM 816872; specimen of indeterminate sex from51°46.9′N177°39.7′E / 51.7817°N 177.6617°E /51.7817; 177.6617 (Onykia robusta specimen (197 cm ML))).Verrill (1876:237) reported a specimen with a mantle length of 232 cm (91.5 in) and a total length of 4.3 m (14 ft) (excluding the ends of the tentacles, which had been destroyed).Nesis (1987:192) likewise gave a maximum mantle length of 230 cm, butRoper & Jereb (2010i:364) wrote that "this old record might be in error", with the species commonly growing to 160 cm ML. Previously known asMoroteuthis robusta (seeBolstad, 2008;Bolstad, 2010).
Megalocranchia maxima185 cm (female)Kubodera & Horikawa (2005:210)Size of female caught offMotobu Peninsula,Okinawa,Japan, identified as "Megalocranchiacf.maxima" (seeKubodera & Horikawa, 2005:223 for photograph). This species is listed under the nameMegalocranchia fisheri in many older sources.Tsuchiya & Okutani (1993),Roper & Jereb (2010c:171) andOkutani (2015a) give maximum of 180 cm, andNorman (2000:158) gives the same forM. fisheri. This species may also be conspecific withMegalocranchia abyssicola (Glaubrecht & Salcedo-Vargas, 2004:65).
Taningia danae
(Dana octopus squid)
170 cmNesis (1982);Roper & Jereb (2010h:266)The largest well documented specimen is a 160 cm ML mature female from the North Atlantic (Roper & Vecchione, 1993:449).
Dosidicus gigas
(Humboldt squid)
150 cmWormuth (1976:38);Norman (2000:165);Glaubrecht & Salcedo-Vargas (2004:54)According toWormuth (1976:38), specimens reaching 150 cm ML are "not uncommon" offPeru.Roperet al. (2010b:301) give a maximum mantle length of 120 cm for specimens offChile and around 100 cm for northern populations, with a more typical mantle length of up to 50–80 cm. The review article ofNigmatullinet al. (2001)—based onc. 230 published papers on the species, in addition to other catch data—also gives a maximum mantle length of 120 cm.
Kondakovia longimana
(giant warty squid)
≈150 cm (estimate; female)Bolstad (2008:171)Estimated size of damaged female (NMV F109447; specimen with 21 mm LRL from63°04.72′S62°56.02′E / 63.07867°S 62.93367°E /-63.07867; 62.93367 (Kondakovia longimana specimen (150 cm ML))).O'Shea (2003b) estimated maximum mantle length as probably exceeding 115 cm. Largest complete specimen measured 108 cm ML (Lynnes & Rodhouse, 2002:1087;Roper & Jereb, 2010i:366).
Mastigoteuthis cordiformis100 cm or moreRoper & Jereb (2010g:253)Based on unpublished reports; largest verified ML is 70 cm (Roper & Jereb, 2010g:253).
Lepidoteuthis grimaldii
(Grimaldi scaled squid)
100 cmRoper & Jereb (2010e:240)
Myopsida (myopsid squids)
SpeciesMaximum mantle lengthReferencesNotes
Loligo forbesi
(long finned squid)
93.7 cmMaximum mantle length is 93.7 cm in males and 46.2 cm in females.[154][155]
Octopoda (octopuses)
SpeciesMaximum mantle lengthReferencesNotes
Haliphron atlanticus
(seven-arm octopus)
69 cm (female)O'Shea (2002:1);O'Shea (2004a:9);Finn (2014a:227)Mantle length of incomplete 2.90 m TL mature female, measured defrosted and wet, prior to fixing. Isolated beaks of comparable size to that of this specimen were recorded byClarke (1986:247–248). Thesexually dimorphic males reach a mantle length of over 10 cm (Finn, 2014a:227).
Enteroctopus dofleini
(giant Pacific octopus)
at least 60 cmNorman (2000:214);Normanet al. (2014:124)
Sepiida (cuttlefish)
SpeciesMaximum mantle lengthReferencesNotes
Sepia apama
(Australian giant cuttlefish)
50 cmReidet al. (2005:68)
Sepia latimanus
(broadclub cuttlefish)
50 cmReidet al. (2005:92)
Sepia hierredda<50 cmReidet al. (2005:88)
Sepia officinalis
(European common cuttlefish)
49 cmReidet al. (2005:99)
Sepia pharaonis
(pharaoh cuttlefish)
42 cmReidet al. (2005:107)
Sepia lycidas
(kisslip cuttlefish)
38 cmReidet al. (2005:96)
Sepia ramani37.5 cmReidet al. (2005:114)
Vampyromorphida (vampire squid)– single extant species
SpeciesMaximum mantle lengthReferencesNotes
Vampyroteuthis infernalis
(vampire squid)
13 cmNesis (1982);Norman & Finn (2014:269)
Sepiolida (bobtail squids)
SpeciesMaximum mantle lengthReferencesNotes
Austrorossia antillensis9 cmReid & Jereb (2005:192)
Rossia pacifica9 cm (female)Reid & Jereb (2005:185)Males grow to 4.5 cm in mantle length.
Rossia macrosoma8.5 cmReid & Jereb (2005:184)More typically the mantle length is 2.0–6.0 cm.
Neorossia caroli8.3 cm (female)Reid & Jereb (2005:190)Males grow to 5.1 cm in mantle length.
Spirulida (spirula)– single extant species
SpeciesMaximum mantle lengthReferencesNotes
Spirula spirula
(ram's horn squid)
rarely exceeds 4.5 cmReid (2005:211)

Total length

[edit]
'Archie', agiant squid specimen at theNatural History Museum inLondon, measures 8.62 m (28.3 ft) in total length (seeAblett, 2012 for more on this specimen)
A bigfin squid (cf.Magnapinna), one of the longest known cephalopods. This specimen was filmed in October 2000 byDSVAlvin in theGulf of Mexico, at 1,940 m (6,360 ft) depth.
A freshly caught specimen ofOmmastrephes bartramii from northernHawaiian waters. This species grows to a total length of 2 m (6.6 ft).
Large specimens ofEnteroctopus dofleini can exceed 3 m (9.8 ft) in total length.

The longest scientifically documented specimens belong to thegiant squid, with a maximum total length of 14–15 m (46–49 ft) (Roper & Shea, 2013:114). Despite its proportionally shorter tentacles, thecolossal squid may rival the giant squid in total length, but the species's size limits are uncertain because only a handful of mature specimens have been recorded.

Teuthida (squids)
SpeciesMaximum total lengthReferencesNotes
Architeuthis dux
(giant squid)
14–15 m (female)Roper & Shea (2013:114)Based on a 40-year data set of more than 50 specimens,Roper & Shea (2013:114) suggest an average total length at maturity of 11 m and a "rarely encountered maximum length" of 14–15 m. Of the nearly 100 specimens examined byRoper, the largest was "46 feet (14 m) long" (Cerullo & Roper, 2012:22).O'Shea & Bolstad (2008) give a maximum total length of 13 m for females based on the examination of more than 130 specimens, measuredpost mortem and relaxed, as well as beaks recovered fromsperm whales (which do not exceed the size of those found in the largest complete specimens).O'Shea estimated the maximum total length for males at 10 m (O'Shea, 2003a).

Older records of 18 m or more were likely exaggerated by stretching of the longfeeding tentacles or resulted from inadequate measurement methods such as pacing (O'Shea & Bolstad, 2008;Roper & Shea, 2013:113).Paxton (2016a) performed a statistical analysis usingliterature records of giant squid specimens and concluded that "squid with a conservative TL of 20 m would seem likely based on current data", but the study has been heavily criticised by experts in the field (Greshko, 2016).

Mesonychoteuthis hamiltoni
(colossal squid)
approaching 9–10 m (estimate)Roper & Jereb (2010c:173)The maximum confirmed total length for this species is 6 m (Rosaet al., 2017).Two specimens ofM. hamiltoni recovered from the stomachs ofsperm whales between 1956 and 1957 off theSouth Shetland Islands andSouth Orkney Islands—both initially identified asArchiteuthis—reportedly measured around 10 and 12 m, respectively (Sweeney & Roper, 2001:56; seeKorabelnikov, 1959:103 andYukhov, 1974:62). Estimated maximum lengths as great as 12–14 m have appeared in the popular literature (seeAnderton, 2007).
cf.Magnapinna
(bigfin squid)
≈7 m (estimate)Vecchioneet al. (2001a:2505);Vecchioneet al. (2001b);Glaubrecht & Salcedo-Vargas (2004:67);Roper & Jereb (2010f:247)Estimate based on specimen observed by commercialROV operated from the oil-drilling shipMillennium Explorer in January 2000,Mississippi Canyon,Gulf of Mexico (28°37′N88°0′W / 28.617°N 88.000°W /28.617; -88.000 (Bigfin squid observed from ROV, January 2000)), at a depth of 2195 m (Vecchioneet al., 2001b).Bolstad (2003) gives an estimate of at least 8 m TL for the largest observed specimen.
Asperoteuthis acanthoderma5.5 m (+)Tsuchiya & Okutani (1993)Total length of immature specimen measuring 0.45 m ML. Much larger specimens of up to 92 cm ML are known (seeKubodera & Horikawa, 2005:223 for photograph).
Onykia robusta
(robust clubhook squid)
4.3 m (14 ft) (+)Verrill (1876:237)Total length of specimen missing ends of tentacles, with a mantle length of 2.32 m (91.5 in) according toVerrill (1876:237).Glaubrecht & Salcedo-Vargas (2004:66) give maximum total length of 4–6 m. Previously known asMoroteuthis robusta (seeBolstad, 2008;Bolstad, 2010).
Galiteuthis phyllura? over 4 m (estimate)Nesis (1985);Ellis (1998a:149);Glaubrecht & Salcedo-Vargas (2004:65)Estimate based on 0.40 m longarm and 1.15 mtentacle from theSea of Okhotsk.[nb 14]Roper & Jereb (2010c:165) cast doubt on the validity of this record.
Dosidicus gigas
(Humboldt squid)
possibly up to 3.7 m (12 ft)Clarke (1966:117);Glaubrecht & Salcedo-Vargas (2004:59)Specimens from theNorthern Hemisphere are much smaller, with those off theCalifornian coast reaching total lengths of less than 1.7 m (Glaubrecht & Salcedo-Vargas, 2004:59).Roperet al. (2010b:301) give maximum total length of close to 2.5 m for specimens offChile.
Megalocranchia maxima2.7 m (+) (female)Young & Mangold (2010)Total length of 1.65 m ML female taken offHawaii (seeYoung & Mangold, 2010 for photograph). Larger specimens of up to 1.85 m ML have been recorded, and these clearly exceed 2.7 m TL (seeKubodera & Horikawa, 2005:223 for photograph).
Taningia danae
(Dana octopus squid)
2.3 m (female)Roper & Vecchione (1993:444)Total length of mature female measuring 160 cm in mantle length, taken from frozen specimen.
Kondakovia longimana
(giant warty squid)
2.25 m (+)Lynnes & Rodhouse (2002:1087)Size of largest complete specimen (1.08 m ML), found floating at surface offSouth Orkney Islands (see alsoCarrington, 2000). Much larger specimens up to an estimated 1.5 m ML are known (Bolstad, 2008:171).
Octopoda (octopuses)
SpeciesMaximum total lengthReferencesNotes
Enteroctopus dofleini
(giant Pacific octopus)
>6.1 mCosgrove (1987)Normanet al. (2014:124) give the maximum total length as "more than 3 m". Questionable length records of up to 9.8 m can be found in the literature (seeHigh, 1976:18).
Cirrinagen. et sp. indet.over 4 m (estimate)Voss (1988);Vecchioneet al. (2008)Estimate based on photographic record; finned octopods are known with certainty to reach at least 1.7 m in total length (Collinset al., 2001).
Haliphron atlanticus
(seven-arm octopus)
4 m (estimate; female)O'Shea (2004a:9);Finn (2014a:227)Estimate based on incomplete 2.90 m mature female, measured defrosted and wet, prior to fixing. Isolated beaks of comparable size to that of this specimen were recorded byClarke (1986:247–248). Males are estimated to reach a total length of 21 cm (Finn, 2014a:227) and perhaps as much as 30 cm (Normanet al., 2002:733).
Vampyromorphida (vampire squid)– single extant species
SpeciesMaximum total lengthReferencesNotes
Vampyroteuthis infernalis
(vampire squid)
≈30 cmNorman & Finn (2014:269)

Mass

[edit]
The 495-kilogram (1,091 lb) colossal squid on display at theMuseum of New Zealand Te Papa Tongarewa. It is the most massive extant cephalopod on record.
The largest documented specimen ofT. danae, preserved at theNational Museum of Natural History. It originally weighed 161.4 kg (356 lb) and had a mantle length of 160 cm (5.2 ft).
A very large mature female ofOnykia robsoni with a mantle length of 88.5 cm (2.90 ft) and total weight of 11.1 kg (24 lb)
Lepidoteuthis grimaldii female measuring 61.7 cm (2.02 ft) in mantle length and weighing 4.07 kg (9.0 lb), from theChatham Rise offNew Zealand (note the prominentdermal cushions, likely used forbuoyancy)
The maximum weight of thegiant Pacific octopus (Enteroctopus dofleini) has been much debated, with numerous reports of specimens exceeding 100 kg (220 lb)
Sepia latimanus is the second largest cuttlefish species, closely rivallingS. apama in both mantle length and mass.

The heaviest known cephalopod, and the largest living invertebrate, is thecolossal squid. The largest recorded specimen of this species, caught in theRoss Sea in 2007, weighed 495 kg (1,091 lb). However, its beak is not the largest known from this species; even bigger colossal squid beaks have been recovered from the stomachs ofsperm whales, indicating that this species can grow larger still.

Teuthida (squids)
SpeciesMaximum massReferencesNotes
Mesonychoteuthis hamiltoni
(colossal squid)
495 kg (female)[Anonymous] (N.d.);Guerra & Segonzac (2014:65);Rosaet al. (2017)Weight of mature female specimen caught in February 2007, measured after thawing. This specimen was originally estimated to weigh 450 kg (Anderton, 2007). Several other specimens with weights in the hundreds of kilograms have been recorded. Beaks recovered fromsperm whale stomachs indicate the existence of even larger specimens, perhaps weighing as much as 600–700 kg ([Anonymous],N.d.).
Architeuthis dux
(giant squid)
275 kg (female)O'Shea (2003a);Grist & Jackson (2007:386);Guerra & Segonzac (2014:65)Maximum size based on the examination of some 105 specimens byO'Shea (2003a), as well as beaks recovered fromsperm whales (which do not exceed the size of those found in the largest complete specimens). Maximum weight for males has been estimated at 150 kg (O'Shea, 2003a; see alsoGrist & Jackson, 2007:386), though heavier specimens have occasionally been reported (seeDeagleet al., 2005 for 190 kg specimen,Hofilena, 2014 for 163 kg specimen).Roper & Jereb (2010a:121) give a maximum weight of up to 500 kg, and "possibly greater". Discredited weights of as much as atonne or more are not uncommon in older literature (O'Shea & Bolstad, 2008).
Taningia danae
(Dana octopus squid)
161.4 kg (female)Roper & Jereb (2010h:266)Weight of 160 cm ML mature female from North Atlantic. Specimen weighed prior to freezing (Roper & Vecchione, 1993:444). According toRoper & Jereb (2010h:266), the previously reported maximum weight of 61.4 kg (based on the same specimen) stems from a typographical error in the original paper ofRoper & Vecchione (1993). This lower value was repeated by a number of subsequent authors, includingSantoset al. (2001:355) andKuboderaet al. (2006:1029).
Onykia robusta
(robust clubhook squid)
50 kgRoperet al. (1984);Roper & Jereb (2010i:364);Guerra & Segonzac (2014:65);Okutani (2015b)Previously known asMoroteuthis robusta (seeBolstad, 2008;Bolstad, 2010).
Dosidicus gigas
(Humboldt squid)
50 kgNigmatullinet al. (2001:10);Roperet al. (2010b:301);[Anonymous] (N.d.)Commonly reaches a maximum weight of around 20–30 kg (Roperet al., 2010b:301). In their introduction to the familyOmmastrephidae,Roperet al. (2010b:269) give a maximum weight of 55–65 kg, but this is contradicted later in the same work by the 50 kg figure in the main species account.Guerra & Segonzac (2014:65) give a maximum weight of 90 kg.
Kondakovia longimana
(giant warty squid)
50 kg (estimate)Guerra & Segonzac (2014:65)Largest complete specimen, found floating at surface offSouth Orkney Islands, had a wet weight of 29 kg and a mantle length of 108 cm (Lynnes & Rodhouse, 2002:1087; see alsoCarrington, 2000), but a damaged female specimen with an estimated mantle length of around 150 cm is known (Bolstad, 2008:171).
Thysanoteuthis rhombus
(diamondback squid)
30 kgMiyaharaet al. (2006);Roper & Jereb (2010k:385)Probably exceeds the recorded mass of 30 kg according toRoper & Jereb (2010k:385).
Ommastrephes bartramii
(neon flying squid)
20–25 kg (female)Roperet al. (2010b:296)Maximum size of specimens from North Atlantic andSouthern Hemisphere (where males reach 2–2.2 kg). Females from the North Pacific are smaller (6 kg), but males may be larger (2–2.9 kg) (Roperet al., 2010b:296).
Megalocranchia maxima23.9 kg (52.7 lb) (+)[Anonymous] (2015b);Mosendz (2015)Mass of specimen found floating dead offHawaii, estimated to measure at least 2.1 m (7 ft) TL, tentatively identified asMegalocranchia fisheri. Larger specimens reaching and even exceeding 2.7 m TL have been recorded, though were apparently not weighed (seeKubodera & Horikawa, 2005:223;Young & Mangold, 2010).
Onykia robsoni
(rugose hooked squid)
11.1 kg (female)Vecchioneet al. (2011)Weight of mature female (88.5 cm ML) caught in bottom trawl at 685–700 m depth overChatham Rise (44°21′S175°32′E / 44.350°S 175.533°E /-44.350; 175.533 (Onykia robsoni specimen (88.5 cm ML))). Previously known asMoroteuthis robsoni (seeBolstad, 2008;Bolstad, 2010).
Octopoda (octopuses)
SpeciesMaximum massReferencesNotes
Enteroctopus dofleini
(giant Pacific octopus)
? >180 kgNormanet al. (2014:124)Cosgrove (1987) andCosgrove & McDaniel (2009:69) gave a maximum confirmed weight of 71 kg for a live specimen collected in the mid-1960s (McClainet al., 2015). There exists a dubious record of a 272 kg specimen which is sometimes cited as the largest known (seeHigh, 1976:18;Hochberg & Fields, 1980:436;Lewy, 2002:65), although it was never actually collected and weighed (Newman, 1994:66;McClainet al., 2015).Normanet al. (2014:124) accept a maximum weight of at least 180 kg, which approximates the 182.3 kg reported for a specimen caught offSanta Barbara,California, in 1945, of which photographic evidence survives (Cosgrove & McDaniel, 2009:67–69). No specimens approaching this size have been reported since the middle of the 20th century, with recent specimens very rarely exceeding 50 kg (Cosgrove & McDaniel, 2009:71). It is possible that the maximum size of the species has decreased over this period, perhaps due tobioaccumulation oftoxicants (Anderson, 2003:3;Cosgrove & McDaniel, 2009:71).
Haliphron atlanticus
(seven-arm octopus)
75 kg (estimate; female)O'Shea (2004a:9)Estimate based on incomplete 61.0 kg mature female, measured defrosted and wet, prior to fixing. Isolated beaks of comparable size to that of the present specimen were recorded byClarke (1986:247–248).
Sepiida (cuttlefish)
SpeciesMaximum massReferencesNotes
Sepia apama
(Australian giant cuttlefish)
>10.5 kgReidet al. (2005:68)
Sepia latimanus
(broadclub cuttlefish)
10 kgReidet al. (2005:92)
Sepia hierredda>7.5 kgReidet al. (2005:88)
Sepia lycidas
(kisslip cuttlefish)
5 kgReidet al. (2005:96)
Sepia pharaonis
(pharaoh cuttlefish)
5 kgReidet al. (2005:107)
Sepia officinalis
(European common cuttlefish)
4 kgReidet al. (2005:99)

Shell diameter

[edit]
Eggcases of six extantArgonauta species (not to scale)
A giant specimen ofArgonauta hians fromTaiwan, which—at roughly 121.5 mm (4.78 in) in diameter—approximates the length of the official world record size shell.
YoungA. argo female with eggcase
Nautilus shells:N. macromphalus (left),A. scrobiculatus (centre), andN. pompilius
Nautilus belauensis offPalau
Two examples of the small internal shell ofSpirula spirula

Nautiluses are the only extantcephalopods with a true external shell; in other groups the shell has been internalised or lost completely. Internal shells include thecuttlebones of cuttlefish, thegladii of squids and the vampire squid, thewinged shells of cirrate octopods, and the spiral shells ofSpirula. Additionally, females of the octopus genusArgonauta secrete a specialised paper-thin eggcase in which they reside, and this is popularly regarded as a "shell", although it is not attached to the body of the animal (seeFinn, 2013).

Cephalopod shell diameter is of interest toteuthologists andconchologists alike. TheRegistry of World Record Size Shells, the most comprehensive publication on maximum shell size in marine molluscs, specifies that specimens "should be measured withvernier type calipers and should reflect the greatest measurable dimension of the shell in any direction including any processes of hard shell material produced by the animal (i.e. spines, wings, keels, siphonal canals, etc.) and not including attachments,barnacles,coralline algae, or any other encrusting organisms" (Pisor, 2008:14). Unlike most other measures of cephalopod size, shell diameter can be determined with a high degree of precision and usually leaves little room for ambiguity. For this reason it is usually recorded to the nearest one-tenth of a millimetre (0.0039 in), as is standard inconchology.

When theRegistry of World Record Size Shells changed ownership in 2008 it was launched as an online database in addition to its print publication. Subsequent rule changes meant that all records required photographic verification. Over time, older records for which photographic evidence could not be obtained were removed from the database. As a result, some records from older editions of the registry actually exceed the size of the current official record holders, sometimes by considerable margins. Where this has occurred, the largest recorded size across all editions is shown first and any discrepancies or competing records are noted thereafter. Where a reliable literature record surpasses all specimens ever included in the registry, this is given instead and the registry record(s) noted thereafter.Pisor (2008) was the fifth and final print edition of the registry published prior to the rule change, andBarbieret al. (N.d.) is the current, continuously updated online database. The registry only covers the shells ofnautiluses andSpirula and the eggcases ofArgonauta.

Octopoda (octopuses)– all extantArgonauta species listed
SpeciesMaximum shell diameterReferencesNotes
Argonauta argo
(greater argonaut)
300.0 mmPisor (2008:22);Barbieret al. (N.d.)Pisor (2008:22) listed this size for a specimen fromAustralia (registered in 1991; in collection ofSIO).Barbieret al. (N.d.) list exactly the same size for a specimen from thePhilippines (year given as 1985; in private collection of Beals Marty). For a discussion of maximum shell size in this species, seeFinn (2013:199). The largest literature record (and the only one to exceed the official world record size) appears to be the rough upper limit of 12 in (304.8 mm) given byWebb (1945:13) for the species as a whole (Finn, 2013:199).

UnderArgonauta cygnus,Pisor (2008:22) listed a 220.0 mm record for a specimen from westernMexico (acquired in 1995; in collection ofSIO). The same specimen was previously listed underArgonauta pacificus (asA. pacifica) byPisor (2005:12). BothA. cygnus andA. pacificus are considered synonyms ofA. argo byFinn (2013) andFinn (2014b).

Argonauta nodosus
(knobbed argonaut)
292.0 mmPisor (2008:22)Size of a specimen fromSouth Australia (registered in 1977; in collection ofAMNH).Barbieret al. (N.d.) list a record of 204.0 mm for a specimen from the "Tasmania area",Australia (year given as 2000; in private collection of B. Henke). For a discussion of maximum shell size in this species, seeFinn (2013:167); a number of unconfirmed reports put the maximum size beyond 300 mm.
Argonauta hians
(muddy argonaut)
121.61 mmBarbieret al. (N.d.)Size of a specimen from theGulf of Aqaba,Red Sea (year given as 1995; in private collection of Simon Weigmann).Pisor (2008:22) listed a record of 112.6 mm for a specimen from thePhilippines (registered in 1988; in private collection of Victor Dan). For a discussion of maximum shell size in this species, seeFinn (2013:186); none of the records discussed therein exceed the current world record specimen.
Argonauta boettgeri[nb 15]108.03 mmBarbieret al. (N.d.)Size of a specimen taken from theChina Sea offZhejiang Province by a local fisherman (year given as 2005; in private collection of Simon Weigmann).Pisor (2008:22) listed a record of 102.2 mm for a specimen from thePhilippines (collected in 2005; in collection of Havelet Marine).Pisor (2005:12) listed a record of 67.0 mm for a specimen fromMozambique, which is thelocus classicus of this species (registered in 2003; in private collection of Pete Stimpson).
Argonauta cornutus[nb 15]98.7 mmBarbieret al. (N.d.)Size of a specimen from westernMexico (year given as 1985; in collection of W. D. Schroeder).Pisor (2008:22) listed possibly the same specimen, also from westernMexico, at 98.6 mm (collected in 1999; in private collection of W. D. Schroeder).
Argonauta nouryi95.5 mmPisor (2008:22);Barbieret al. (N.d.)Size of a specimen fromCalifornia (collected in 1992; in collection of Havelet Marine). For a discussion of maximum shell size in this species, seeFinn (2013:207); none of the records discussed therein exceed the current world record specimen (when includingA. cornutus, which Finn treats as a synonym ofA. nouryi).
Nautilida (nautiluses)– all extant species listed
SpeciesMaximum shell diameterReferencesNotes
Nautilus pompilius pompilius
(emperor nautilus)
254.0 mmPisor (2008:121);Barbieret al. (N.d.)Size of a specimen fromIndonesia (registered in 2003; in private collection of Pete Stimpson), listed asN. p. pompilius.Hutsellet al. (1997:48) listed a 253.0 mm specimen, also fromIndonesia (collected in 1983; in private collection of Cecelia Abbott).Harasewych & Moretzsohn (2010:632) give a maximum shell diameter of 268 mm for this species, but this is based on an erroneous record.[nb 16]

UnderN. repertus (which is treated here in synonymy withN. p. pompilius;Jereb (2005:53) considered it a "questionable species"),Pisor (2005:93) listed a 230.0 mm record for a specimen fromIndonesia (registered in 2000; in private collection of Pete Stimpson),Pisor (2008) did not include the taxon at all, whileBarbieret al. (N.d.) list a record of 242.07 mm for a specimen fromIndia (no year given; in private collection of Simon Weigmann).

The largestalbinisticN. p. pompilius is listed byBarbieret al. (N.d.) as a 175.0 mm specimen trawled at 400 m depth offZamboanga in thePhilippines (year given as 2013; in collection of Havelet Marine).

Nautilus stenomphalus
(white-patch nautilus)
239.39 mmBarbieret al. (N.d.)Size of a specimen fromTimor Island,Indonesia (year given as 2009; in private collection of Simon Weigmann). UnderN. pompiliusf.stenomphalus,Pisor (2008:121) listed a maximum shell diameter of 221.0 mm for a specimen from thePhilippines (no year given; in collection of Havelet Marine).
Nautilus belauensis
(Palau nautilus)
239 mmGrulke (2016:76)Grulke (2016:76) gives an adult shell size range of 180–239 mm, and a mean adult shell diameter of 200 mm.Jereb (2005:54) gives 226 mm as the maximum size for the species, with no reference to a particular specimen.Pisor (2008:121) andBarbieret al. (N.d.) list a record of 221.0 mm for a specimen fromBabeldaob,Palau (year given as 1980; in collection of Havelet Marine).
Nautilus pompilius suluensis220.0 mmBarbieret al. (N.d.)Size of a specimen from thePhilippines (year given as 2000; in private collection of Pete Stimpson).Pisor (2008:121) listed a maximum shell diameter of 148.0 mm for a specimen from thePhilippines (registered in 2000; in private collection of Pete Stimpson).N. p. suluensis is a dwarf form from theSulu Sea that has the smallest mean shell diameter of all known extant nautilus populations, at 115.6 mm (Dunstanet al., 2011).
Allonautilus scrobiculatus
(crusty nautilus)
215.0 mmPisor (2005:93)Size of a specimen from thePhilippines (registered in 2000; in private collection of Pete Stimpson), listed asNautilus scrobiculatus.Pisor (2008) did not include this species at all.Barbieret al. (N.d.) list a record of 214.0 mm for a specimen fromIndonesia (year given as 2013; in private collection of Pete Stimpson).
Nautilus macromphalus
(bellybutton nautilus)
180.62 mmBarbieret al. (N.d.)Size of a specimen fromNew Caledonia (year given as 2008; in private collection of Simon Weigmann).Pisor (2008:121) listed a maximum shell diameter of 180.0 mm for a specimen from New Caledonia (collected in 1995; in private collection of Kent Trego).
Allonautilus perforatusaround 180 mmJereb (2005:55);Grulke (2016:83)Given as the maximum size for the species, with no reference to a particular specimen.Jereb (2005:55) consideredA. perforatus a "[v]ery rare form of questionable validity".
Spirulida (spirula)– single extant species
SpeciesMaximum shell diameterReferencesNotes
Spirula spirula
(ram's horn squid)
28.8 mm[Anonymous] (2003c)Size of a specimen fromSamar Island, thePhilippines (collected in 2003).Pisor (2008:139) andBarbieret al. (N.d.) list a record of 27.2 mm for a specimen fromPhuket Island,Thailand (collectedc. 2000; in collection of Havelet Marine; see[Anonymous], 2006 for online record).

Extinct taxa

[edit]
The largestammonite ever found, a specimen ofParapuzosia seppenradensis measuring 1.742 m (5.72 ft) in diameter. The fossil pictured is the original held at theWestfälisches Museum für Naturkunde inMünster, but many casts exist.
The earliest discovered fossil ofP. seppenradensis, as it appeared at theWestphalian Museum of Natural History in 1895. A human skeleton provided scale, while an oil painted background reconstructed the living chamber and animal itself, together with assortedCretaceous sealife.
The largest ammonite ever found inBulgaria: a 1.44 mParapuzosia sp. collected nearShumen in 1932. It is on display at theNational Museum of Natural History inSofia.
Giant fossil ofPachydesmoceras pachydiscoide fromMikasa,Hokkaido,Japan, exhibited at theNational Museum of Nature and Science inTokyo
Titanites giganteus at theNatural History Museum inLondon
The uncoiled shell ofBaculites grandis, a species ofheteromorph ammonite
Fossilised guards of theJurassic belemnitesMegateuthis gigantea (top two) andMegateuthis aalensis
Fossilised rostrum ofAcroteuthis subquadrata
Endoceras orthocones
Artist's reconstruction ofOrthoceras sp., anorthocerid of theMiddle Ordovician
Ammonoidea (ammonoids)
SpeciesMaximum shell diameter (length for heteromorphs)ReferencesNotes
Parapuzosia seppenradensis2–3.5 m (estimate)Landois (1895:100);Landois (1898:27);Teichert & Kummel (1960:6);Summesberger (1979:128);Holland (1987:6);Kennedy & Kaplan (1995:21);Lewy (2002:66); frimet al. (2021)Widely recognised as the largest ammonite specimen ever found (Payneet al., 2009:27;Grulke, 2014:124). Discovered in 1895 in a quarry inSeppenrade,Coesfeld,Germany, the original is on display at theWestfälisches Museum für Naturkunde inMünster. Estimate based onlectotype measuring 1.742 m in diameter (Kennedy & Kaplan, 1995:21) with an incompleteliving chamber, assuming living chamber took up one-fourth of the outerwhorl.Teichert & Kummel (1960:6) suggested an even larger shell diameter of around 3.5 m for this specimen, assuming the body chamber extended for three-fourths to one full whorl.Landois (1898:27) estimated the total live weight at 1456 kg, of which the shell would constitute 705 kg. The fossil itself weighs around 3.5 tonnes (Beer, 2015). A smaller specimen of 1.36 m was found in the same quarry some years earlier (Beer, 2015). In 1971 a portion of an ammonite possibly surpassing the Seppenrade specimen was reportedly found in abrickyard inBottrop, western Germany (Beer, 2015). In 2021, diameter of 2 m is estimated for the largest specimen.[156]
Parapuzosia bradyi>1.8 m (estimate)Larsonet al. (1997:44);Lewy (2002:66)Largest known North American ammonite. Estimate based on incomplete specimen measuring 1.37 m in diameter (missing at least half awhorl of thebody chamber).
Peltoceratinaegen. et sp. indet.1.78 m (estimate)Poulton (1989)Estimate based on small portion ofouter whorl measuring 1.2 m along the venter and subtending a chord of 1.13 m. The estimate is based on the ultimate whorl height/diameter ratio of"Titanites"occidentalis (about 35%), and assumes a constant rate of expansion. More crude calculations give a circular diameter of 2–2.4 m (best fit of the specimen's outline to a curve yields 2.16 m estimate).
Eopachydiscus sp.1.67 mGrulke (2014:125)This specimen, from theAlbianDuck Creek Formation ofTexas, has been exhibited at the Tucson Fossil Show and in a New York auction.
Pachydesmoceras cf.pachydiscoide1.65 m (estimate)Kin & Niedźwiedzki (2012:19)Estimate based on 0.98 m diameter specimen representing an apparently completephragmocone (previously referred toLewesiceras peramplum orParapuzosia). A more complete and therefore larger specimen (1.18 m diameter) consisting of a complete phragmocone and near-completebody chamber is also known (Kin & Niedźwiedzki, 2012:17).
Lytoceras taharoaense1.5 mStevens (1978a:3);Stevens (1978b:2);Stevens (1979a:33);Stevens (1979b:6);Stevens (1979c:166);Stevens (1985:153);Grulke (2014:126)Size based on essentially complete shell with only some damage to theaperture.
Mesopuzosia mobergi<1.5 mKin & Niedźwiedzki (2012:19)
Parapuzosia austeni<1.5 mKin & Niedźwiedzki (2012:19)Puzosia mayoriana is a synonym.
Moutoniceras sp.1.47 m [heteromorph]Grulke (2014:126)In private collection of Wolfgang Grulke, according to whom it is likely the largestheteromorph ammonite ever found (Grulke, 2014:126). Originating fromMorocco it is displayed in part of the original rock matrix withsympatricGassendiceras heteromorphs. Its unrolled shell length would have exceeded 3 m (Grulke, 2014:126). A specimen claimed to be very slightly larger at 1.48 m was auctioned in September 2015 ([Anonymous], 2015a).
Parapuzosia bosei1.45 mScott & Moore (1928:276);Lewy (2002:66)From theAustin Chalk of theRio Grande region,Texas. The largest known specimen was reported byScott & Moore (1928:273) to be 4 ft 9 in (1.45 m) in diameter and "impossible to extract from its matrix". The authors found "[m]any others only slightly smaller", of which three were collected in 1928 and deposited atTexas Christian University (Scott & Moore, 1928:273–274).
Parapuzosia americana1.37 m (estimate)Scott & Moore (1928:276)From theAustin Chalk of theRio Grande region,Texas.
Corbinites occidentalis1.37 mFrebold (1957:66);Westermann (1966)Originally assigned to the genusTitanites.[157] Size based on specimen consisting of animprint and part of the lastwhorl preserved as aninternal mould.
Diplomoceras maximum>1 m [heteromorph]Olivero & Zinsmeister (1989)
Tropaeum imperatoralmost 1 mGrulke (2014:126)Largest ammonite known fromAustralia.Grulke (2014:126) writes: "No exact size is available but it could be almost 1 m across".
Belemnoidea (belemnoids)
SpeciesMaximum rostrum measurementsReferencesNotes
Megateuthis sp.0.7 mTL (Dvms: 30 mm;Dvma: 50 mm)Schlegelmilch (1998:1);Weis & Mariotti (2007:166);Ibaet al. (2015:23–24)Megateuthis elliptica is "the longest belemnite species known", with rostra from theHumphriesianum Zone inRumelange andLuxembourg reaching 60–70 cm (Weis & Mariotti, 2007:166). The whole animal is estimated to have been 3–5 m long (Eyden, 2003a). Fuchs and Stinnesbeck (2021) estimated total mantle length of 2.22 m.[158]
Belemnitinagen. et sp. indet.?TL (Dvms: 30 mm;Dvma: ?)Ibaet al. (2015:23)Known from a single incomplete rostrum (TCSM-J1-0001) from thePliensbachianTeradani Formation inTeradani,Toyama Prefecture,Japan. The specimen is missing the apical and alveolar regions and comprises only the middle (stem) region of the rostrum. It measures 45 mm in total length by 30 mm and 25 mm across at the anterior and posterior ends, respectively.Ibaet al. (2015:23) wrote: "In theBelemnitina, the diameter of the alveolar region is generally larger than those of the apical and stem regions. Thus maximum rostrum diameter of the Teradani specimen is estimated to reach much more than 30 mm."
Acroteuthis sp.?TL (Dvms: 39 mm;Dvma: 42 mm)Ibaet al. (2015:23)One of "the largest belemnites ever observed", with a rostrum comparable to that of the indeterminatebelemnitinid from Teradani.
Pachyteuthis sp.?TL (Dvms: 39 mm;Dvma: 40 mm)Ibaet al. (2015:23)One of "the largest belemnites ever observed", with a rostrum comparable to that of the indeterminatebelemnitinid from Teradani.
Belemnitinagen. et sp. indet.?TL (Dvms: ?;Dvma: >33 mm)Ibaet al. (2015:23)From theHettangianNiranohama Formation of northeastern Japan. One of "the largest belemnites ever observed", with a rostrum comparable to but likely slightly smaller than that of the indeterminatebelemnitinid from Teradani.
Nautiloidea (nautiloids)
SpeciesMaximum shell lengthReferencesNotes
Endoceras giganteum5.733 m (estimate)Kluget al. (2015:270)Estimate based on incomplete 3-metre-long shell deposited at theMuseum of Comparative Zoology,Harvard University.Teichert & Kummel (1960:5) assumed length of 8 m based onbody chamber-to-phragmocone ratio of 1:2 and wrote that this was likely "the largest fragment of an endoceroid cephalopod on display anywhere in the world". However,Kluget al. (2015:270) estimated the total length of the complete shell at 5.733 m, with a volume of 158.6 litres. From theKatian ofNew York (Kluget al., 2015:270). The published literature includes unconfirmed reports of even larger endocerids.Frey (1995) mentioned length of 6 m as maximum size for genusCameroceras, although this author support theory that some large specimens ofEndoceras belong toCameroceras instead.
Endoceras proteiforme3.0–4.6 m (10–15 ft)Clarke (1897:778);Teichert & Kummel (1960:1)Size based on "entire shells" (Clarke, 1897:778).
Rayonnoceras solidiforme2.8 m (estimate)Kluget al. (2015:270)From theVisean ofArkansas. Shell volume estimated at 62.5 litres.
Deiroceras hollardi2.6 m (estimate)Kluget al. (2015:270)From the earlyEmsian of "Jebel Mdouar". Shell volume estimated at 68.3 litres.
Cameroceras turrisoidesc. 2 m (estimate)Kröger & Ebbestad (2014)From theKatian ofSweden. Estimated after maximum diameter (165 mm).[159][160]
Actinoceridagen. et sp. indet.1.911 m (estimate)Kluget al. (2015:270)From theLlandovery ofGotland. Shell volume estimated at 8.9 litres.
Orthoceridagen. et sp. indet.1.783 m (estimate)Kluget al. (2015:270)From theLudlow ofGotland. Shell volume estimated at 4.1 litres.
Ormoceras TUG 1308-11.72 m (estimate)Kluget al. (2015:270)From theSandbian ofEstonia. Shell volume estimated at 2.7 litres.
Ormoceras giganteum MB.C.119401.71 m (estimate)Kluget al. (2015:270)From theDarriwilian. Shell volume estimated at 2.7 litres.
Lambeoceras lambii1.405 m (estimate)Leith (1942:130);Teichert & Kummel (1960:4)Estimate based on incomplete 1.155 m long shell.
Orthoceras regarium1.39 m (estimate)Kluget al. (2015:270)From theWenlock of Joachimsthal.[nb 17] Shell volume estimated at 5.1 litres.
Temperoceras aequinudum1.333 m (estimate)Kluget al. (2015:270)From theLochkovian of "Ouidane Chebbi". Shell volume estimated at 9.2 litres.
Zeravshanoceras priscum1.299 m (estimate)Kluget al. (2015:270)From theEifelian. Shell volume estimated at 1.6 litres.
Ordogeisonoceras amplicameratum>1.25 mFrey (1995:40)Shell diameter up to 10.5 cm. Originally described asOrthoceras amplicameratum.Orthoceras ludlowense is considered a synonym.
Cameroceras hennepini<1.2 m (4 ft) (estimate)Clarke (1897:779)Size estimate based on "the most complete of the fragments which represent it".
Actinoceras vaughanianum1.198 m (estimate)Kluget al. (2015:270)From theSerpukhovian ofOklahoma. Shell volume estimated at 8.7 litres.
Polygrammoceras? cf.P. sp. A1.13 m (estimate)Frey (1995:69)Estimate based on a "single, very large fragment of a phragmocone". Shell diameter to 9.0 cm.
Plagiostomoceras sp.1.1 m (estimate)Kluget al. (2015:270)From theGivetian ofOnondaga, New York. Shell volume estimated at 0.0052 litres.
Endoceras decorahense1.06 m (estimate; phragmocone only)Miller & Kummel (1944);Teichert & Kummel (1960:2)Size estimate based on two portions of aninternal mould of thephragmocone, measuring 62.5 cm and 32 cm, with an estimated missing middle section of 11.5 cm.
Proterovaginoceras incognitum1 m (estimate)Kluget al. (2015:270)From theDapingian ofJämtland,Sweden. Shell volume estimated at 0.8 litres.
Teuthida (squids)
SpeciesMaximum mantle lengthReferencesNotes
Yezoteuthis giganteus≈1.7 m (estimate)Tanabeet al. (2006:142)Size estimate based on preservedupper jaw measuring 97.0 mm in maximum length, similar to that of the largestgiant squid (Architeuthis dux).Tanabeet al. (2006:143) wrote that this species "appears to be the largest fossil coleoid ever described".
Octopodiformes
SpeciesMaximum mantle lengthReferencesNotes
Enchoteuthis melanaeup to 2 m (estimate)Eyden (2003b); Fuchset al. (2020)Total length estimated to be up to 3 m.Tusoteuthis longa is likely to be invalid, and largest specimen should belongs toE. melanae.[161]
Leptotheuthis1.3 m (gladius length)Fuchs and Stinnesbeck (2021)Reference spells "Leptotheuthis".[158]
Eromangateuthis soniae1.2 m (gladius length)Fuchs (2019)Size based ongladius fromQueensland,Australia.[162] A second gladius measuring more than a metre and showing possible evidence of predation byKronosaurus is also known (Eyden, 2003b).
Trachyteuthis0.8 m (gladius length)Fuchs and Stinnesbeck (2021)Size given for Early Cretaceous species.[158]
Nanaimoteuthis hikidai≈0.7 m (estimate)Tanabeet al. (2015)Estimated from hood length of lower jaw with proportion of vampire squid.[96] Authors also estimated mantle length from fossil vampyromorphs unofficially, resulting ≈1.6 m mantle length.[163]
Nanaimoteuthis yokotai≈0.54 m (estimate)Tanabeet al. (2010)Estimated from hood length of lower jaw with proportion of vampire squid.[164]
Sister group ofdecapodiformes
SpeciesMaximum mantle lengthReferencesNotes
Longibelus matsumotoi (?)91.1-98.9 cm (estimate)Tanabe and Misaki (2023)Estimated from rostral length of lower jaw, with proportion ofSpirula, giant squid and colossal squid.[97]

Anatomical superlatives

[edit]

Eyes

[edit]
Preserved giant squid eye
Gonatus sp. swimming past a submersible during a dive to theKermadec Arc offNew Zealand.Gonatids such as this have proportionately slightly larger eyes than do giant squid.[165]

Thegiant andcolossal squids have the largest recordedeyes of any living animal, with a maximum diameter of at least 27 cm (11 in) and a 9 cm (3.5 in)pupil.[5] This is three times the size of the largestfish eyes—up to 90 mm (3.5 in) inswordfish—and more than twice the diameter of the largestwhale eyes—up to 109 mm (4.3 in), 61 mm (2.4 in), and 55 mm (2.2 in) inblue,humpback, andsperm whales, respectively—which are the largest amongvertebrates.[5] A large colossal squid caught in 2014 and dissected at theMuseum of New Zealand Te Papa Tongarewa had eyes some 35–37 cm (14–15 in) across.[166] There are unconfirmed reports from the 19th century of giant squid eyes up to 40 cm (1.3 ft) across.[167] Only the extinctichthyosaurs are known to have approached these dimensions,[168] with some species having eyes up to 35 cm (14 in) in diameter.[169]

Despite their size, the eyes of giant and colossal squids do not appear to be disproportionately large; they do not deviate significantly from theallometric relationship seen across other squid species.[170]Sepiolids are noted for having exceptionally large eyes, which are much bigger relative to their mantle length than those of the giant squid; the same is true ofHistioteuthis species.[165]Gonatids and theloliginidsLoligo andLolliguncula also have proportionately somewhat larger eyes thanArchiteuthis.[165] Some sources state that thevampire squid (Vampyroteuthis infernalis) has the largest eyes of any animal relative to its size, with a 15 cm (5.9 in) specimen having eyes around 2.5 cm (0.98 in) in diameter.[171]

Iridoteuthis iris is an extreme example of asepiolid with disproportionately large eyes

There is some debate in the scientific community as to the evolutionary reason behind the extremely large eyes of giant and colossal squids.[172]Nilssonet al. (2012) andNilssonet al. (2013) argue that it is ananti-predator adaptation for enhanced detection ofsperm whales, with the squids picking upplanktonbioluminescence triggered by moving whales, perhaps from distances exceeding 120 m (390 ft).Schmitzet al. (2013a) andSchmitzet al. (2013b) contend that their eyes are so large due to aphylogenetically conserved developmental pattern that governs the relative dimensions of squids and their eyes, and that anyfitness benefits their size may confer in terms of predator avoidance are the result ofexaptation ("pre-adaptation").

Neurons

[edit]

Squid giant axons can exceed 1 mm (0.039 in) in diameter: 100 to 1000 times the thickness ofmammalianaxons. The axons of theHumboldt squid (Dosidicus gigas) are exceptional in that they can reach a diameter of as much as 1.5 mm (0.059 in), and those ofLoligo forbesii can also exceed 1 mm.[173] Such was the importance of Humboldt squid toelectrophysiology research that when the animals migrated out of reach ofChilean fishermen in the 1970s "it led to the demise of a world-class electrophysiology laboratory" based there.[174] Squid giant axon diameters do not necessarily correlate with overall body size; those of thegiant squid (Architeuthis dux) are only 0.137–0.21 mm (0.0054–0.0083 in) thick.[173]

Diagram showing the three major elements (red, green, and yellow) of the squid giant neuronal system. The arrows indicate the direction of transmission from the head ganglion towards the mantle. The funnel (light blue) is the site of rapid water expulsion following mantle contraction.

Thesquid giant synapse is the largestchemical junction in nature. It lies in the stellateganglion on each side of the midline, at the posterior wall of the squid's muscular mantle. Activation of this synapse triggers a synchronous contraction of the mantle musculature, causing the forceful ejection of a jet of water from the mantle. Thiswater propulsion allows the squid to move rapidly through the water and, in the case of the so-called 'flying squids', even to jump through the surface of the water (breaking the air–water barrier) to escape predators.[175] Many essential elements of how all chemical synapses function were first discovered by studying the squid giant synapse.[176]

Photophores

[edit]

Taningia danae, a very largeoctopoteuthid squid, possesses "lemon-sized" yellowphotophores at the tips of two of itsarms, which are the largest known light-emitting organs in the animal kingdom.[177] Video footage shot in 2005 in deep water offJapan showsT. danae emitting blindingflashes of light from these photophores as it attacks its prey.[178] A pair of muscular lids surrounds each photophore and it is the withdrawal of these lids that produces the flashes. A large individual filmed from a remote submersible offHawaii in 2015 can clearly be seen opening the lids to reveal its photophores.[179] It is believed that this highly manoeuvrable squid uses bright flashes to disorientate potential prey. The flashes may also serve to illuminate prey for easier capture or play a role in courtship and/or territorial displays.[180]


Reproductive organs

[edit]
A dissected male specimen ofOnykia ingens, showing an erect 67 cm (2.20 ft) penis approximating the combined length of the mantle, head, and arms

Extreme penis elongation has been observed in the deep water squidOnykia ingens. When erect, the penis may be as long as the mantle, head, and arms combined.[181] As such, deep water squids have the greatest known penis length relative to body size of all mobile animals, second in the entire animal kingdom only to certain sessilebarnacles.[182]

See also

[edit]

Notes

[edit]
  1. ^The heaviest scientifically validated cephalopod specimen was acolossal squid (Mesonychoteuthis hamiltoni) weighing 495 kg (1,091 lb),[1] while hatchlings ofIllex illecebrosus—some of the smallest known—have a mass of around 0.00015 g (5.3×10−6 oz).[2] The mass ratio between the two is(495,0000.00015)=3,300,000,000{\displaystyle \left({\frac {495,000}{0.00015}}\right)=3,300,000,000}.
    Comparing only adult females, the difference between the colossal squid and the pygmy squidIdiosepius thailandicus—which weighs 0.20 g (0.0071 oz) at maturity[3]—would be(495,0000.2)=2,475,000{\displaystyle \left({\frac {495,000}{0.2}}\right)=2,475,000}.
  2. ^Wood & O'Dor (2000:93) elaborated on this mass estimate as follows:

    There are [...] no published weights of hatchlingNautilus spp. The weight of a hatchlingN. belauensis was estimated using hatchling shell size and a regression analysis of the cubed shell diameter versus the weight of seven youngN. belauensis that weighed <50 g [1.8 oz] [...] in addition to a single hatchlingN. pompilius that was weighed for the present study on 24 April 1996 at theWaikiki Aquarium. The hatchlingN. pompilius weighing 4.33 g [0.153 oz], with a maximum shell diameter of 26.25 mm [1.033 in], fit a highly significant correlation [...] between cubed shell diameter and weight, which indicates that a hatchlingN. belauensis with a 30 mm [1.2 in] shell diameter [...] would weigh approximately 5.9 g [0.21 oz].

  3. ^Normanet al. (2002:733) wrote: "The most extreme examples of sexual size dimorphism come from marine or parasitic taxa where females are difficult to locate (Ghiselin 1974)."
  4. ^In particular,Scott Cassell mentioned a giantHumboldt squid he named 'Scar' (after the scars on its mantle), whose size he described thus:

    Scar is over seven feet [2.1 m] in length, larger than anyone has ever proven. His body was 3 feet [0.9 m] thick (He's too big to put my arms around him) and I estimated he weight [sic] between 230 and 250 pounds [100–110 kg].[75]

  5. ^Depending on how one defines 'shell-bearing', other extinct contenders could include the aquatic turtlesArchelon andStupendemys (the former having a skeletal framework rather than a solid shell), the tortoiseMegalochelys, and thearmoured mammalGlyptodon and its relatives.
  6. ^Iwai (1956:139) reported on two small squid (92 and 104 mm ML) recovered from the "digestive canal" of asperm whale, which he identified as belonging to the genusArchiteuthis.Roper & Young (1972:220) showed that this was certainly a misidentification and attributed them instead to the familyPsychroteuthidae. In a brief summary of this case,Ellis (1998a:121) gave an erroneous total length of "8 feet" (2.4 m) for the larger of the two specimens (even though in an almost word-for-word identicalfootnote inEllis, 1994a:145 he correctly stated the total length was "8 inches" [20 cm]). This mistake was repeated byGlaubrecht & Salcedo-Vargas (2004:67), giving rise to the claim of an implausibly large psychroteuthid "with about three meter total length".
  7. ^ab
    T. W. Kirk's sketch of theArchiteuthis longimanustype specimen in lateral aspect, fromKirk (1888). Note the extreme length of the feeding tentacles relative to the mantle and arms.
    Thomas William Kirk gave the following account of this specimen in hisformal description ofArchiteuthis longimanus:

    [...] Early last month Mr. Smith, a local fisherman, brought to the[Colonial] Museum the beak and buccal-mass of a cuttle which had that morning been found lying on the "Big Beach" (Lyall Bay), and he assured us that the creature measured sixty-two feet [18.9 m] in total length. I that afternoon proceeded to the spot and made a careful examination, took notes, measurements, and also obtained a sketch, which, although the terribly heavy rain and driving southerly wind rendered it impossible to do justice to the subject, will, I trust, convey to you some idea of the general outline of this most recently-arrived Devil-fish.

    Measurements showed that, although Mr. Smith was over the mark in giving the total length as 62 feet [18.9 m] (probably, not having a measure with him, he only stepped the distance), those figures were not so very far out; for, although the body was in all ways smaller than any of the hitherto-described New Zealand species, the enormous development of the very slight tentacular arms brought the total length up to 55 feet 2 inches [16.81 m], or more than half as long again as the largest species yet recorded from these seas.
    The length of the tentacular arms is not a very important character, as they are known to be capable of extension or retraction at the will of the animal, at least to a considerable extent.

    [...] This specimen was a female, and to this fact may be due some of the points in which it differs from previous occurrences; but yet they are so considerable that I have no doubt a new subgenus at least will have to be created for its reception. In the meantime I place it underArchiteuthis, with the full knowledge, however, that it cannot possibly remain there permanently, the shape of the arms and the fins alone being sufficient to put it out of association with that genus. As soon as opportunity offers, I hope to make a further study and fully determine its affinities.[112]

    Kirk (1888:38) provides a table with a detailed breakdown of the specimen's various measurements. There is, however, a discrepancy between the total length of 684 in (17.37 m, or exactly 57 ft) given in the table—which agrees with the individual values of 71 in (1.80 m) for the mantle, 22 in (0.56 m) for the head, and 591 in (15.01 m) for the tentacles—and the total length of 55 ft 2 in (16.81 m) given by Kirk in the body of the article.

    Wood (1982:191) suggested that, due to the tentacles' highly retractile nature, the total length of 62 feet (18.9 m) originally reported by the fisherman "may have been correct at the time he found the squid", and that "[t]his probably also explains the discrepancy in Kirk's figures". Owing to its small mantle size,Wood (1982:191) estimated that "this specimen probably weighed less than 300 lb [140 kg]".

    O'Shea & Bolstad (2008) opined that the reported total length of 55 ft 2 in (16.81 m) "simply cannot be correct" and attributed it to either "imagination" or artificial lengthening of the tentacles. They added that a female giant squid with a mantle length of 71 in (180 cm) "measured post mortem and relaxed (by modern standards) today would have a total length of ≈32 feet [9.8 m]".

  8. ^abKirk (1880:312) wrote of this specimen:

    On 23rd of May last, theVen.Archdeacon Stock very kindly sent me word that three boys, named Edward R. Stock, and Frank and Walter Morrah, had that morning discovered, atLyall Bay, what they took to be a very large cuttlefish, with arms several feet long. I lost no time in proceeding to the spot, determined, if possible, to carry home the entire specimen; but judge my surprise when, on reaching the bay, I saw an animal of the size represented in the drawing now before you.*Victor Hugo'saccount of his "pieuvre" was brought vividly to my mind, and I could not help thinking that a man would stand but a poor chance if he once got within the grasp of such a monster.

    My first step after spreading out the arms, was to make a rough sketch and very careful measurements. I then proceeded to extract the so-calledskeleton, but found that some person or persons, who had visited the spot earlier than myself, had not been able to resist the temptation to try the temper of their knives upon its back, and had in consequence severed thecuttle-bone in various places. However, I was able, not only to procure all the pieces, but also thebeak,tongue, and some of the suckers, only a few of which remained, the greater portion of them having been torn off, either in some fierce encounter, or during the rough weather which had prevailed for some days previously.
    The length of body from tip oftail to anterior margin of the mantle was 9 feet 2 inches [2.79 m] and 7 feet 3 inches [2.21 m] in circumference; the head from anterior margin of mantle to roots of arms 1-foot 11 inches [58 cm], making the total length of the body 11 feet 1 inch [3.38 m]. The head measured 4 feet [1.2 m] in circumference. The sessile arms measured 4 feet 3 inches [1.30 m] in length, and 11 inches [28 cm] in circumference; each of these arms bore thirty-six suckers, arranged in two equal rows (as shown by the scars), and measuring from1316{\displaystyle {\tfrac {13}{16}}} to14{\displaystyle {\tfrac {1}{4}}} of an inch [20.6–6.4 mm] in diameter; every sucker was strengthened by a bony ring armed with from forty to sixty sharp incurved teeth. The tentacular arms had been torn off at the length of 6 feet 2 inches [1.88 m], which was probably less than half their original length.
    The fins were posterior, and were mere lateral expansions of the mantle, they did not extend over the back as in the case withOnychoteuthis, etc.; each measured 24 inches [61 cm] in length and 13 inches [33 cm] in width.

    Thecuttle bone, when first extracted, measured 6 feet 3 inches [1.91 m] in length, and 11 inches [28 cm] in width, but has since shrunk considerably; it was broadly lanceolate, with a hollow conical apex 118{\displaystyle {\tfrac {1}{8}}} inch [28.6 mm] deep.

  9. ^abThe Thimble Tickle specimen, whichAddison Emery Verrill referred to as his specimen No. 18,[113] is often cited as the largest recorded giant squid, and has long been treated as such byGuinness.[114]The Thimble Tickle squid was found aground offshore, alive, on 2 November 1878, near Little Bay Copper Mine,Thimble Tickle,Notre Dame Bay,Newfoundland. It was secured to a tree with agrapnel and rope and died as the tide receded. No part of the animal was retained as it was cut up for dog food. The length of the "body [..] from the beak to the extremity of the tail" (i.e. mantle plus head) was said to be 20 ft (6.1 m), with "one of the arms" (presumably a tentacle) measuring 35 ft (10.7 m), for a total length of 55 ft (16.8 m).[113] ReverendMoses Harvey detailed the encounter in a letter to theBoston Traveller dated 30 January 1879, which was reproduced byVerrill (1880a:191–192):

    On the 2d day of November last, Stephen Sherring, a fisherman residing in Thimble Tickle, not far from the locality where the other devil-fish [Verrill specimen No. 19; seeVerrill, 1880a:192], was cast ashore, was out in a boat with two other men; not far from the shore they observed some bulky object, and, supposing it might be part of a wreck, they rowed toward it, and, to their horror, found themselves close to a huge fish, having large glassy eyes, which was making desperate efforts to escape, and churning the water into foam by the motion of its immense arms and tail. It was aground and the tide was ebbing. From the funnel at the back of its head it was ejecting large volumes of water, this being its method of moving backward, the force of the stream, by the reaction of the surrounding medium, driving it in the required direction. At times the water from the siphon was black as ink.

    Finding the monster partially disabled, the fishermen plucked up courage and ventured near enough to throw the grapnel of their boat, the sharp flukes of which, having barbed points, sunk into the soft body. To the grapnel they had attached a stout rope which they had carried ashore and tied to a tree, so as to prevent the fish from going out with the tide. It was a happy thought, for the devil fish found himself effectually moored to the shore. His struggles were terrific as he flung his ten arms about in dying agony. The fishermen took care to keep a respectful distance from the long tentacles, which ever and anon darted out like great tongues from the central mass. At length it became exhausted, and as the water receded it expired.

    The fishermen, alas! knowing no better, proceeded to convert it into dog's meat. It was a splendid specimen—the largest yet taken—the body measuring 20 feet [6.1 m] from the beak to the extremity of the tail. It was thus exactly double the size of the New York specimen [Verrill specimen No. 14; seeVerrill, 1880a:189], and five feet [1.5 m] longer than the one taken by [fisherman William] Budgell [No. 19]. The circumference of the body is not stated, but one of the arms measured 35 feet [10.7 m]. This must have been a tentacle.

    Such is the fame of the Thimble Tickle specimen that a Giant Squid Interpretation Centre and a "life-sized", 55-foot (17 m) sculpture have been built near the site of its capture.[115] The sculpture appeared on aCanadian postage stamp issued in 2011.[116]

  10. ^
    Giant squid beak being measured. Isolated beaks found in the stomachs of predators such as sperm whales can be used to estimate the size of the original animal.[122]
    Romanovet al. (2017) used the following allometric scaling equations to estimate the dorsal mantle length (DML) from thelower rostral length (LRL) of the beak, and total length (TL) from DML:


    LRL=11.2×log10DML19.3{\displaystyle LRL=11.2\times log_{10}DML-19.3} (all values in mm)[123]
    DML=e(1.168+98.031×LRL){\displaystyle DML=e^{(-1.168+98.031\times LRL)}} (all values in m)[124]
    TL=0.006+5.117DML{\displaystyle TL=0.006+5.117DML} (all values in m)[124]
    TL=1.59×DML1.98{\displaystyle TL=1.59\times DML^{1.98}} (all values in m)[125]
  11. ^Aldrich restated this belief in "Monsters of the Deep", the second episode of the 1980 television seriesArthur C. Clarke's Mysterious World: "I believe the giant squid reach an approximate maximum size of something like one hundred and fifty feet [46 m]".[135]Richard Ellis, apparently unaware of Aldrich's similar statements in print, commented: "It is difficult to imagine why Aldrich would have made such an irresponsible statement, unless it had to do with being on camera".[136]Arthur C. Clarke himself once remarked: "it would be strange indeed if the world's biggest squid had been among the very few cast ashore to be examined and measured by naturalists. There may well be specimens more than a hundred feet [30 m] in length."[137]
  12. ^An example from the May 2000 issue ofDiver magazine reads: "The generally accepted record was of awesome proportions, weighing 272kg and with an arm span of 9.6m."[149]
  13. ^Wood (1982:191) provided the following details: "DrAnna M Bidder (pers. comm.) of the Department of Zoology atCambridge University, possesses a transverse slice of thepen of anotherMesonychoteuthis which, judging by its width, must have come from acranchid [sic] measuring at least 5 m [16 ft] in mantle length." The same information is summarised byBright (1989:146).
  14. ^ab
    Drawing ofGaliteuthis phyllura from theBulletin of the United States Fish Commission (1881), showing the proportionately short length of the arms and tentacles
    Ellis (1998a:148–149) wrote of this specimen:

    The Russian vesselNovoulianovsk, working in theSea of Okhotsk in 1984, brought up the remains of a gigantic specimen ofGaliteuthis phyllura from a depth of one thousand to thirteen hundred meters (thirty-three hundred to forty-three hundred feet), andNesis (1985) said that it was "almost as large asMesonychoteuthis hamiltoni (of the same family)." Only an arm and a tentacle were collected, but they were so large (the arm was 40 cm long [15.6 inches] and the tentacle 115 cm [45.5 in]) that Nesis was able to estimate the mantle length at 265 to 275 cm (8.69 to 9.02 ft), and the total length at over 4 meters (more than 13 feet). "Because of its narrow body," wrote Nesis, "we conclude that its mass is consistently lower than that of the other large squids."

    Roper & Jereb (2010c:165) questioned the validity of this record, writing: "this is considered a doubtful record that might refer to total length; probably the maximum mantle length is less than 400 to 500 mm". ButSteve O'Shea commented:

    The late, greatKir Nesis was not one to exaggerate, and must have had very good reason to cite a possible mantle length of 2.7m [8.9 ft] forGaliteuthis phyllura. Not only would such a monstrous mantle render this the 'largest species ofGaliteuthis', but it would give the species a mantle that was considerably longer than that ofArchiteuthis, and marginally longer (0.2m [0.66 ft]) than the submature Colossal Squid (Mesonychoteuthis) we reported last year [in 2003].
    What amazes me is that the adult has never made the press (that I am aware of). Even ifGaliteuthis lacks hooks on the arms it would still be a most frightening squid to bump into in the abyss. (O'Shea, 2004b)

  15. ^abThe taxonomic validity of theseArgonauta species is questionable.Finn (2013) andFinn (2014b) recognised only four species in the genus:A. argo (syn.A. cygnus,A. pacificus),A. hians (syn.A. boettgeri),A. nodosus, andA. nouryi (syn.A. cornutus).
  16. ^This record is based on a shell collected in 2001 from theTimor Sea offIndonesia, which was sold in 2003 as a specimen ofN. repertus with a diameter of 268 mm (10.6 in). It was subsequently found to measure only around 243 mm (9.6 in) and the discrepancy was put down to an encoding mistake ([Anonymous], 2003a). Another specimen from the same locality, sold around the same time, was claimed to measure 255 mm (10.0 in) ([Anonymous], 2003b).
  17. ^There are two places which bear or have borne the name Joachimsthal:Jáchymov, Czech Republic andJoachimsthal, Brandenburg, Germany. The citation quotes an unpublished report, and it is unclear which one it is referring to. If the report is later than 1945, when the official name of the Czech town was changed, it is most likely the place in Germany.

References

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Short citations

[edit]
  1. ^Rosaet al., 2017:1871
  2. ^abcWood & O'Dor, 2000:93
  3. ^abcNabhitabhata, 1998:28
  4. ^O'Dor & Hoar, 2000:8
  5. ^abcNilssonet al., 2012:683
  6. ^Smithet al., 2016
  7. ^abTeichert & Kummel, 1960:6
  8. ^abcVermeij, 2016
  9. ^Pierceet al., 1995;2004
  10. ^Gerhardt, 1966:171;Muntz, 1995;Ellis, 1998a:11
  11. ^Salvador & Tomotani, 2014
  12. ^Ley, 1941
  13. ^Lee, 1883;Ellis, 1994b
  14. ^seeGarcin & Raynal, 2011;Barrère, 2017
  15. ^Guerraet al., 2011
  16. ^Tratz, 1973;Ellis, 1997a,b
  17. ^Landman & Ellis, 1998;Ablett, 2012
  18. ^abRoper & Voss, 1983:58
  19. ^Ellis, 1998a:106;Glaubrecht & Salcedo-Vargas, 2004:62
  20. ^Roper & Voss, 1983:55
  21. ^seeArkhipkinet al., 2015
  22. ^abcO'Shea & Bolstad, 2008
  23. ^Roper & Young, 1972:205
  24. ^Clarke, 1996:1105;O'Shea & Bolstad, 2008
  25. ^seeClarke, 1962;Wolff, 1981;1984;Grögeret al., 2000;Staudingeret al., 2009;Potieret al., 2011
  26. ^Clarke, 1986:11
  27. ^Zeidberg, 2004:4195;Phillips, 2004:ii
  28. ^Nabhitabhata, 1998:32
  29. ^O'Doret al., 1986:59;Wood & O'Dor, 2000:93
  30. ^Cosgrove & McDaniel, 2009:88
  31. ^Grulke, 2014:105
  32. ^Rowlett, Joe (2017-10-06)."Meet The World's Smallest & Weirdest Squid,Idiosepius". Reefs.com. Retrieved19 January 2019.
  33. ^Boletzky, 2003:19
  34. ^Boletzky, 1999:24;2003:20;Normanet al., 2002:733
  35. ^Normanet al., 2002:733;Fairbairn, 2007:3
  36. ^Normanet al., 2002:733
  37. ^seeKennedy & Cobban, 1990
  38. ^Nishiguchi, M.K., R. Mapes (2008)."Cephalopoda"(PDF).University of California Press. pp. 163–199.{{cite web}}: CS1 maint: multiple names: authors list (link)
  39. ^Rosa & Seibel, 2010;McClainet al., 2015;Rosaet al., 2017
  40. ^abcdefgMcClainet al., 2015
  41. ^Collinset al., 2001;Collins & Villanueva, 2006:294
  42. ^Voss, 1988;Vecchioneet al., 2008
  43. ^seeTeichert & Kummel, 1960;Stevens, 1988;Eyden, 2003b;Larson, 2010;Ibaet al., 2015
  44. ^O'Shea & Bolstad, 2008;[Anonymous],N.d.
  45. ^ab[Anonymous],N.d.
  46. ^Lovis, 2011
  47. ^[Anonymous],N.d.
  48. ^[Anonymous],N.d.
  49. ^abc[Anonymous],N.d.
  50. ^O'Shea, 2003c
  51. ^Ellis, 1998b;Paxton, 2016a;Bittel, 2016;Romanovet al., 2017
  52. ^abPaxton, 2016a
  53. ^abKuboderaet al., 2016
  54. ^seeHolroyd, 2005
  55. ^Cerullo & Roper, 2012:22
  56. ^abO'Shea, 2003a
  57. ^abO'Shea & Bolstad, 2008;Roper & Shea, 2013:113
  58. ^Bittel, 2016
  59. ^abGreshko, 2016
  60. ^Deagleet al., 2005
  61. ^Hofilena, 2014
  62. ^[Anonymous], 2000;Wong, 2000
  63. ^Vecchioneet al., 2010
  64. ^seeBolstad, 2008;2010
  65. ^Norman, 2000:174;Bolstad, 2008:107;Okutani, 2015b
  66. ^Roperet al., 1984;Roper & Jereb, 2010i:364;Okutani, 2015b
  67. ^seeDall, 1873;Verrill, 1876;Phillips, 1933;Croker, 1934;Phillips, 1961;Smith, 1963
  68. ^Hoff, 2003:90
  69. ^Nigmatullinet al., 2001;Roperet al., 2010b:301
  70. ^Wormuth, 1976:38;Norman, 2000:165;Glaubrecht & Salcedo-Vargas, 2004:54
  71. ^abcRoperet al., 2010b:301
  72. ^Clarke, 1966:117;Glaubrecht & Salcedo-Vargas, 2004:59
  73. ^Glaubrecht & Salcedo-Vargas, 2004:59
  74. ^Nigmatullinet al., 2001:10;Roperet al., 2010b:301
  75. ^abCassell, 2005
  76. ^abLynnes & Rodhouse, 2002:1087; see alsoCarrington, 2000
  77. ^Bolstad, 2008:171
  78. ^Guerra & Segonzac, 2014:65
  79. ^Cosgrove, 1987
  80. ^Norman, 2000:214;Normanet al., 2014:124
  81. ^abCosgrove & McDaniel, 2009:67–69
  82. ^abCosgrove & McDaniel, 2009:71
  83. ^Anderson, 2003:3;Cosgrove & McDaniel, 2009:71;Yong, 2015
  84. ^O'Shea, 2002:1;2004a:9;Finn, 2014a:227
  85. ^O'Shea, 2004a:9
  86. ^Landois, 1895:108
  87. ^Carnall, 2017
  88. ^Kennedy & Kaplan, 1995:21
  89. ^Landois, 1895:100
  90. ^Beer, 2015
  91. ^[Anonymous],N.d.;[Anonymous], 2008
  92. ^Teichert & Kummel, 1960:2
  93. ^Flower, 1955:329
  94. ^Landois, 1898:27
  95. ^Grulke, 2014:124
  96. ^abTanabe, Kazushige; Misaki, Akihiro; Ubukata, Takao (2014)."Late Cretaceous record of large soft-bodied coleoids based on lower jaw remains from Hokkaido, Japan".Acta Palaeontologica Polonica.doi:10.4202/app.00052.2013.ISSN 0567-7920.
  97. ^abTanabe, Kazushige; Misaki, Akihiro (2023-06-20)."Upper Cretaceous record of non-belemnitid coleoid jaws from Hokkaido, Japan, and its evolutionary implications".Cretaceous Research.151 105624.Bibcode:2023CrRes.15105624T.doi:10.1016/j.cretres.2023.105624.ISSN 0195-6671.S2CID 259228292.
  98. ^abcdEllis, 1998a:211
  99. ^seeEllis, 1998a;Salcedo-Vargas, 1999;Glaubrecht & Salcedo-Vargas, 2004
  100. ^Robson, 1933:681; see for exampleMitsukuri & Ikeda, 1895;Meek & Goddard, 1926;Clarke & Robson, 1929;Phillips, 1933;Croker, 1934;Rees, 1950;Smith, 1963;Nesis, 1970
  101. ^Ellis, 1998a:211;Norman, 2000:174
  102. ^Poppe & Goto, 1993
  103. ^Kubodera & Mori, 2005;Kubodera, 2010:25
  104. ^[Anonymous], 2007;Kubodera, 2010:38
  105. ^[Anonymous], 2013;[NHK], 2013c
  106. ^Baird, 2002
  107. ^[Anonymous], 2002a;O'Shea, 2003d
  108. ^e.g.Guerraet al., 2018
  109. ^Coe, 1929:36;G.E. Verrill, 1958:69; seeVerrill 1874a,b;1875a,b,c;1876;1877;1878;1880a,b;1881a,b;1882a
  110. ^O'Shea & Bolstad, 2008;Dery, 2013;Yong, 2015
  111. ^seeHansford, 2009
  112. ^Kirk, 1888:35–36
  113. ^abVerrill, 1880a:191;Verrill, 1880b:285
  114. ^seeWood, 1982:189;Carwardine, 1995:240;Glenday, 2014:62
  115. ^Hickey, 2009
  116. ^Hickey, 2010;[Anonymous],N.d.
  117. ^Paxton, 2016a:86
  118. ^Murray, 1874:121
  119. ^abcdRomanovet al., 2017
  120. ^Paxton, 2016a:83
  121. ^seeMurray, 1874;Verrill, 1875a;Verrill, 1875b
  122. ^seeRoeleveld, 2000;Cherel, 2003;Romanovet al., 2017
  123. ^Roeleveld, 2000:185
  124. ^abPaxton, 2016a:85
  125. ^McClainet al., 2015:Table 2
  126. ^seeSanderson, 1956;Heuvelmans, 1958;Ley, 1959
  127. ^Roper & Boss, 1982:97
  128. ^Wood, 1982:193
  129. ^Roper & Boss, 1982:99;Wood, 1982:192;Haszprunar & Wanninger, 2012:R510
  130. ^Ellis, 1998a:143
  131. ^seeEllis, 1998a:142
  132. ^Matthews, 1938
  133. ^Ley, 1959:210
  134. ^abAldrich, 1980:59
  135. ^Ellis, 1998a:6–7
  136. ^Ellis, 1998a:7
  137. ^Ellis, 1998a:6
  138. ^Ellis, 1998a:142
  139. ^seeFeldman, 1999 for more on this specimen
  140. ^O'Shea, 2003a;Roper & Jereb, 2010a:121
  141. ^Alexander, 1998:1233
  142. ^abEllis, 1998a:106
  143. ^MacGinitie & MacGinitie, 1949
  144. ^MacGinitie & MacGinitie, 1968;Wood, 1982:190
  145. ^Heuvelmans, 1958
  146. ^Ellis, 1998a:107
  147. ^Dall, 1885:432
  148. ^Newman, 1994:66
  149. ^Bavendam, 2000:63–64
  150. ^Newman, 1994:66;Cosgrove & McDaniel, 2009:66–67
  151. ^Anderson, 2003:2;Cosgrove & McDaniel, 2009:71
  152. ^Anderson, 2003:3;Cosgrove & McDaniel, 2009:71;Scheel & Anderson, 2012
  153. ^Anderson, 2003:2
  154. ^The Azorean Loligo forbesi (Cephalopoda: Loliginidae) in captivity: transport, handling, maintenance, tagging and survival
  155. ^Body size and fin length as determinants in the geographic distribution of Loliginid squids
  156. ^Ifrim, Christina; Stinnesbeck, Wolfgang; González, Arturo H. González; Schorndorf, Nils; Gale, Andrew S. (2021-11-10)."Ontogeny, evolution and palaeogeographic distribution of the world's largest ammonite Parapuzosia (P.) seppenradensis (Landois, 1895)".PLOS ONE.16 (11) e0258510.Bibcode:2021PLoSO..1658510I.doi:10.1371/journal.pone.0258510.ISSN 1932-6203.PMC 8580234.PMID 34758037.
  157. ^Poulton, Terence P. (2023)."Corbinites (Subfamily Lithacoceratinae), a new genus for the giant western Canadian Late Kimmeridgian or Tithonian (Late Jurassic) ammonite Titanites occidentalis Frebold".Volumina Jurassica.21:27–38.ISSN 1731-3708.
  158. ^abcFuchs, Dirk; Stinnesbeck, Wolfgang (2021-11-01)."Large-sized gladius-bearing octobrachians (coleoid cephalopods) in the Turonian plattenkalk of Vallecillo, Mexico".Cretaceous Research.127 104949.Bibcode:2021CrRes.12704949F.doi:10.1016/j.cretres.2021.104949.ISSN 0195-6671.
  159. ^Kröger, Björn (2013-03-27)."The cephalopods of the Boda Limestone, Late Ordovician, of Dalarna, Sweden".European Journal of Taxonomy (41).doi:10.5852/ejt.2013.41.ISSN 2118-9773.
  160. ^Kröger, Björn; Ebbestad, Jan Ove R. (2014). "Palaeoecology and palaeogeography of Late Ordovician (Katian–Hirnantian) cephalopods of the Boda Limestone, Siljan district, Sweden".Lethaia.47 (1):15–30.Bibcode:2014Letha..47...15K.doi:10.1111/let.12034.ISSN 0024-1164.
  161. ^Fuchs, Dirk; Iba, Yasuhiro; Heyng, Alexander; Iijima, Masaya; Klug, Christian; Larson, Neal L.; Schweigert, Günter (2020). Brayard, Arnaud (ed.)."The Muensterelloidea: phylogeny and character evolution of Mesozoic stem octopods".Papers in Palaeontology.6 (1):31–92.Bibcode:2020PPal....6...31F.doi:10.1002/spp2.1254.ISSN 2056-2802.S2CID 198256507.
  162. ^Fuchs, Dirk (2019-10-08)."Eromangateuthis n. gen., a new genus for a late Albian gladius-bearing giant octobrachian (Cephalopoda: Coleoidea)".Paleontological Contributions.doi:10.17161/1808.29619.hdl:1808/29619.ISSN 1946-0279.
  163. ^Kitakyushu Museum of Natural History & Human History (2015-03-05)."史上最大級のイカ類とタコ類の化石発見".
  164. ^Tanabe, Kazushige; Hikida, Yoshinori (2010). "Jaws of a New Species ofNanaimoteuthis(Coleoidea: Vampyromorphida) from the Tuironian of Hokkaido, Japan".Paleontological Research.14 (2):145–150.Bibcode:2010PalRe..14..145T.doi:10.2517/1342-8144-14.2.145.ISSN 1342-8144.S2CID 128429338.
  165. ^abcSchmitzet al., 2013a
  166. ^Farquhar, 2014;[Te Papa], 2014
  167. ^Land & Nilsson, 2012:86
  168. ^Motaniet al., 1999;Humphries & Ruxton, 2002
  169. ^Nilssonet al., 2012:687
  170. ^Schmitzet al., 2013a:45
  171. ^Ellis, 1996:177;[Anonymous], 2005a; though seeYounget al., 2015
  172. ^Partridge, 2012
  173. ^abAdelman & Gilbert, 1990:102
  174. ^Scully, 2008
  175. ^Arata, 1954;Murata, 1988;Maciáet al., 2004;Muramatsuet al., 2013;O'Doret al., 2013
  176. ^seeLlinás, 1999
  177. ^Ellis, 1998a:149;Barrat, 2015
  178. ^seeKuboderaet al., 2006
  179. ^seeBarrat, 2015
  180. ^Kuboderaet al., 2006:1033
  181. ^Arkhipkin & Laptikhovsky, 2010:299;Walker, 2010
  182. ^Arkhipkin & Laptikhovsky, 2010:300

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