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Home , Abraliopsis, Ancistrocheirus, Argonaut (animal), Austrorossia, Brachioteuthidae, Brachioteuthis

PHYLUM: : CEPHALOPODA

Authors

Rob Leslie1 and Marek Lipinski2

Citation

Leslie RW and Lipinski MR. 2018. Phylum Mollusca – Class Cephalopoda In: Atkinson LJ and Sink KJ (eds) Field Guide to the Ofshore Marine Invertebrates of South , Malachite Marketing and Media, Pretoria, pp. 321-391.

1 South African Department of Agriculture, Forestry and Fisheries, Cape Town 2 Ichthyology Department, Rhodes University, Grahamstown,

321 Phylum: MOLLUSCA Class: Cephalopoda

Argonauts, octopods, cuttlefish and

Introduction to the Class Cephalopoda

Cephalopods are among the most complex and The relative length of the arm pairs, an important advanced invertebrates. They are distinguished from identiication character, is generally expressed as the rest of the Phylum Mollusca by the presence an arm formula, listing the arms from longest to of circumoral (around the mouth) appendages shortest pair: e.g. III≥II>IV>I indicates that the two commonly referred to as arms and . lateral arm pairs (Arms II and III) are of similar length irst appeared in the Upper , and are longer than the ventral pair (Arms IV). The over 500 million years ago, but most of those dorsal pair (Arms I) is the shortest. ancestral lineages went extinct. Only the (Subclass Nautiloidea) survived past the (400 Vampyromorpha (Vampire squids) million years ago) and are today represented by only This order contains a single . Body sac-like, two surviving genera. All other living cephalopods black, gelatinous with one pair (two in juveniles) of belong to the Subclass that irst appeared paddle-like ins on and a pair of large light in the late Palaeozoic (400-350 million years ago). organs at the base of the ins; the eight circumoral appendages have deep webs; a pair of long, thin, Subclass Coleoidea ilamentous appendages that can be retracted into Coleoidea are characterised by possessing eight pits on the outer crown between Arms I and and II; or ten circumoral appendages armed with suckers, arms with a single row of stalked suckers lacking suckers modiied into hooks in some ; chitinous rings, lanked by a row of cirri on either shell internal, reduced or absent. The family-level side. taxa of living cephalopods are well-resolved and accepted. However, although most families can be Order Spirulida (Ram’s horn squids) sorted into groups, there is considerable debate on Ten circumoral appendages; internal shell well- the relationships between, and to a lesser extent developed, spirally coiled and chambered, visible within, these groups – see Jereb and Roper (2005) externally; ins small, positioned on posterior edge for several classiication schemes that have been of mantle. proposed. For isheries purposes, length frequency data are recorded as mantle length (ML; Figures 1-3) Order Sepiida (Cuttleish and bobtail squids) measured in centimetres or millimetres. Ten circumoral appendages (eight arms and two tentacles – Figure 2); tentacles can be retracted Order Octopoda (Octopods) into pockets between Arms III and IV; eyes covered Sac-like body with eight circumoral appendages by a cornea. Cuttleish (): shell straight, armed with sessile suckers (without stalks) without well-developed, calcareous or chitinous; ins long, chitinous rings. Arm pairs are numbered from fringing the dorsal-lateral edge of mantle. Bobtail dorsal to ventral (Figure 1). There are two suborders. squids (): shell rudimentary; ins wide, Suborder : suckers in one or two rows rounded, attached about midway along mantle. without cirri; body irm, well-muscled (all octopods in this guide) or soft and gelatinous; ins absent. The structure of the club (Figure 5), presence or Suborder Cirrata: suckers in a single row lanked by absence of suckers at the tips of the dorsal arms a row of cirri (Figure 4); body soft, semi-gelatinous; and whether the ventral mantle margin is entire or a pair of paddle-like ins. emarginated (Figure 6) are important ield characters for identiication of cuttleish.

322 Phylum: Mollusca Cephalopoda

Mantle length

I

II

III

IV Funnel

Hectocotylised arm (modiied arm for sperm transfer, tip called the ligula)

Figure 1: Schematic of a generalised incirrate

Mantle length

I II III

IV Fin Club Funnel

Figure 2: Schematic of a generalised cuttleish

Mantle length

III Fin II

I IV

Tail

Fin lobe Funnel-locking cartilage

Carpus

Figure 3: Schematic of a generalised

323 Phylum: Mollusca Cephalopoda

Orders and Oegopsida (Squids) the number of buccal lappets (Figure 8), the number Two closely related orders (sometimes treated as and position of on the eyeballs, and suborders). Ten circumoral appendages (eight arms the presence or absence of hooks on the arms and/ and two tentacles – Figure 3); tentacles cannot be or clubs. retracted into pockets, reduced or absent in adults of some species; eyes covered by a membrane, General cornea (Myopsida) or open to seawater (Oegopsida); Distribution maps are based on records in the stalked suckers with chitinous rings (modiied into Research Survey database for surveys conducted hooks in some species); photophores present in between years 1986 and 2016 by the RS Africana, RV many species (on internal organs, externally in Dr Fridtjof Nansen, FV Andromeda and FV Compass mantle, on the eyeballs or on the arms); mantle can Challenger. Records are augmented with specimens be locked to the head and funnel using the nuchal- from Iziko Museum, Cape Town. All photographs, and funnel-locking cartilages respectively (fused to except where noted otherwise, are copyright of RW head and funnel in some species). Leslie.

The shape of the funnel-locking cartilage (Figure 7), Acknowledgements found at the lateral corners of the funnel just under Illustrations from the three-volume work, the ventral mantle margin (Figure 3), is an important Cephalopods of the World (Jereb & Roper 2005, 2010; identiication character. Other important characters Jereb et al., 2014) are used with permission from the are whether the buccal connective is attached to the Food and Agriculture Organization of the United dorsal or ventral edge of the ventral arms (Figure 8), Nations.

a) b) c)

Figure 4: Oral view of typical Cirrate octopod Figure 5: Example cuttleish clubs with a) small showing suckers lanked by cirri subequal suckers, b) moderately enlarged and c) greatly enlarged medial suckers

324 Phylum: Mollusca Cephalopoda

Figure 6: Ventral mantel of cuttleish showing entire (left) and deeply emarginated (right) ventral margin

Figure 7: Examples of shapes of funnel-locking Figure 8: Buccal of squids cartilage

325 Phylum: Mollusca Cephalopoda

Argonauta argo (ArgArg)

Phylum: Mollusca

Class: Cephalopoda

Order: Octopoda

Suborder: Incirrata

Family:

Common: Greater (Paper ) Alternate: -

Lateral ribs aligned Lateral ribs smooth with keel tubercles continuous

Terminal tubercles Tubercles in pairs on paired on either side either side of keel of dorsal keel

Dorsal keel width and tubercle size constant No lateral tubercles

10 mm

Distinguishing features Size • Head small, embedded within mantle. Eyes Females attain 97 mm ML, 300 mm shell length. prominent, protruding and constricted at base. Males 9 mm ML. • present. All arms with two rows of suckers. Webs between arms shallow. Distribution • : Arms I thick at base, length variable, large Circumglobal between 40° N and 40° S. Pelagic, membranous lap extending full length of arm. surface to 200 m on both West and South Coasts. Arms IV more than 3x ML and 20-30% longer than Arms II. Arm formula IV > II > III. Similar species • : Small. 12-13 suckers on normal arms. A. hians and A. nodosus: Shell dorsal keel width • Lateral ribs smooth, continuous or branched and tubercle size increasing with growth, i.e. from from axis to keel, aligned with keel tubercles. towards mouth. Keel tubercles not paired, • Dorsal keel narrow and constant width around alternating on either side of the keel. : Arms circumference of shell. IV shorter than Arms II; with 10-11 (A. hians) or • Keel tubercles consistent in size and arranged 17-20 (A. nodosus) suckers on normal arms. in pairs with a ridge across keel between pairs. References Jereb et al., 2014; Nesis, 1987; Sanchez, 1988. Left Arm III. Long, slender, self-amputating extension (almost as long as the arm) kept coiled in sac below left eye.

326 Phylum: Mollusca Cephalopoda

Argonauta hians (ArgHia)

Phylum: Mollusca

Class: Cephalopoda

Order: Octopoda

Suborder: Incirrata

Family: Argonautidae

Common: Lesser argonaut (Paper nautilus) Alternate: -

Wide dorsal keel Large tubercles alternate on Ribs prominent, widely either side of dorsal keel spaced

Lateral ribs not aligned with keel tubercles

Dorsal keel width and tubercle size increase No lateral with growth Dorsal keel width increases tubercles 10 mm with growth

Distinguishing features Size • Head small, embedded within mantle. Eyes Females attain 40 mm ML, 106 mm shell length. prominent, protruding and constricted at base. Males 7 mm ML. • Ink sac present. All arms with two rows of suckers. Webs between arms shallow. Distribution • : Arms I thick at base, length variable, large Oceanic on both coasts. Pelagic, surface to 200 m membranous lap extending the full length depth. of arm. Arms II & III 1.4x to 2x ML and 20-50% longer than Arms IV. Arm formula III ≥ II > IV. Similar species • : Small. 10-11 suckers on normal arms. A. argo: Lateral ribs smooth; dorsal keel narrow, • Lateral ribs prominent smooth without width and tubercle size constant; keel tubercles tubercles, not aligned with keel tubercles. arranged in pairs. : Arms IV longest (more than • Dorsal keel width and tubercle size increase 3x ML); : 12-13 suckers on arms. with growth (i.e. from apex towards ). Tubercles alternate on either side of keel. A. nodosus: Lateral ribs inconspicuous, ending in a • Dorsal keel wide. chain of separate tubercles; shell white. : Arms II longer than Arms III, 2.0 to 2.8 times ML; : 17-20 Hectocotylus suckers on normal arms. Left Arm III. Long, slender, self-amputating extension (almost as long as the arm) kept coiled in sac below References left eye. Jereb et al., 2014; Nesis, 1987.

327 Phylum: Mollusca Cephalopoda

Argonauta nodosus (ArgNod)

Phylum: Mollusca

Class: Cephalopoda

Order: Octopoda

Suborder: Incirrata

Family: Argonautidae

Common: Knobbed argonaut (Paper nautilus) Alternate: -

Lateral ribs a chain of Tubercles alternate on Tubercles alternate on separate tubercles either side of dorsal either side of keel keel

Dorsal keel width and tubercle size increase with growth Dorsal keel width moderate, 10 mm width increases with growth

Distinguishing features Distribution • Head small, embedded within mantle. Eyes Circumglobal in between prominent, protruding and constricted at base. 10° S and 44° S. Pelagic, surface to 200 m depth. • Ink sac present. All arms with two rows of suckers. Webs between arms shallow. Similar species • : Arms I thick at base, length variable, A. argo: Lateral ribs smooth; dorsal keel narrow, width large membranous lap extending the whole and tubercle size constant; keel tubercles arranged length. Arms II 2.0 to 2.8 times ML; Arms III & IV in pairs. : Arms IV longest (more than 3x ML); subequal. Arm formula II > III ≈ IV. : 12-13 suckers on normal arms. • : Small. 17-20 suckers on normal arms. A. hians: Lateral ribs prominent, smooth, not • Lateral ribs ending in a chain of separate terminating in keel tubercle; dorsal keel 20-30% of tubercles terminating in an acute keel tubercle. shell length. Shell of-white to brown. : Arms II & • Dorsal keel width and tubercle size increase III subequal, 1.4 to 2.0 times ML; : 10-11 suckers with growth (i.e. from apex towards aperture). on normal arms. Tubercles alternate on either side of keel.

Hectocotylus References Jereb et al., 2014; Nesis, 1987. Left Arm III. Long, slender, self-amputating extension (almost as long as the arm) kept coiled in sac below left eye.

Size Females attain 138 mm ML, 292 mm shell length. Males 11 mm ML.

328 Phylum: Mollusca Cephalopoda

Bathypolypus valdiviae (BatVal)

Phylum: Mollusca

Class: Cephalopoda

Order: Octopoda

Suborder: Incirrata

Family: Bathypolypodidae

Common: - Alternate: -

FEMALE MALE

Head narrower than globular body Ligula with four transverse laminae

Arms short, webbed for Small suckers in one-third of length two alternating rows

Hectocotylised arm Hectocotylised 50 mm right arm III 10 mm

Distinguishing features Distribution • Ink sac absent. Both coasts, but more common on West Coast. • Small, smooth, purplish octopod with ovoid Generally 450 to 1000 m depth, but has been muscular mantle. recorded at 200 m. • Head narrower than body, eyes slightly protuberant. Similar species • Interbranchial web pouches absent. Enteroctopus and Octopus: Arms moderate length • A single papilla over each eye. (3.5-5.0 times mantle length); ink sac present. • Arms short, subequal with two rows of small Benthoctopus: Arms three to six times mantle length; suckers, webbed for 33% of length. large prominent suckers; ink sac absent. Arm formula I ≈ II ≈ III ≈ IV. schultzei (inshore, under 20 m depth) and Hectocotylus Velodona togata (KZN): Distinguished by single Right Arm III. Ligula a broad, rounded disc with a row of suckers on arms (to date neither have been deep trough bearing four big transverse laminae. recorded on demersal surveys).

Size References 80 mm mantle length. Jereb et al., 2014; Roper et al., 1984; Nesis, 1987; Sanchez, 1988.

329 Phylum: Mollusca Cephalopoda

Benthoctopus berryi (BenBer)

Phylum: Mollusca

Class: Cephalopoda

Order: Octopoda

Suborder: Incirrata

Family:

Common: Deepwater octopus Alternate: -

Arms 3-6 x mantle length with prominent suckers

Smooth skin on mantle

Suckers in two Buccal area and alternating ventral surface of rows arms dark brown

Arm III hectocotylised, ligula short and narrow, 5-7% arm length

Detail of suckers on arms 50 mm

Distinguishing features Similar species • Ink sac absent. Enteroctopus magnificus: Ink sac present; • Buccal area and ventral surface of arms characteristic fold of loose skin at end of mantle; chocolate brown. lacks the dark pigmentation on the buccal area and • Suckers large, prominent, arranged in two ventral surfaces of arms; Arm formula II = I > III = IV. alternating rows, i.e. not arranged in pairs. • Arms three to six times longer than mantle. Octopus vulgaris: Ink sac present; lacks the dark Arms I, II and III subequal in length and longer pigmentation on the buccal area and ventral than Arms IV. Arm formula: I ≈ II ≈ III > IV. surfaces of arms; Arm formula II = III > I = IV.

Hectocotylus valdiviae: Small, purple, with short Right Arm III. Ligula short, narrow 5-7% of subequal arms; ink sac absent. hectocotylised arm length. Eledone schultzei (inshore, under 20 m depth) and Size Velodona togata (KZN): Distinguished by single 50 mm ML. row of suckers on arms (to date neither have been recorded on demersal surveys). Distribution Rare endemic. West Coast from 600-2 200 m. References Jereb et al., 2014; Nesis, 1987.

330 Phylum: Mollusca Cephalopoda

Enteroctopus magniicus (OctMag)

Phylum: Mollusca

Class: Cephalopoda

Order: Octopoda

Suborder: Incirrata

Family: Octopodidae

Common: Southern giant octopus Alternate: Octopus doleini (in error); Octopus magniicus Records from shallow waters may be the result of misidentiications

Arms similar in length, but dorsal pairs (Arms I & II) slightly longer than ventral pairs (Arms Loose skin III & IV) forms a fold at end of mantle Colour usually has reddish tones

Hectocotylised right Arm III with very long Hectocotylised ligula right Arm III 100 mm (from Jereb et al., 2014)

Distinguishing features Similar species • Ink sac present. Octopus vulgaris: Lateral arms distinctly longer than • Large and robust, without enlarged suckers on medial arms (III ≥II>IV>I); two to three pairs enlarged arms. suckers on lateral arms; generally smaller and found • Arm length moderate 3.5-5.0 times ML; at shallower depths; colour usually greyish rather subequal in length. Arm formula II = I > III = IV. than reddish tones; lacks the loose skin on the • Colour usually with reddish tones; distinctive mantle. Ligula small (only 2.5% of arm length) and fold of loose skin at the end of the mantle. spoon-shaped. • No large erectile papillae on dorsal mantle; Benthoctopus: Ventral surface of arms dark brown; single large papilla and three or four cirri over suckers prominent. Arm formula I = II = III = IV. each eye. Bathypolypus: Small, purple with short arms; ink Hectocotylus sac absent. Right Arm III. Ligula long (16-25% of length of arm), Eledone schultzei (inshore, under 20 m depth) and tapering to a blunt terminus. Velodona togata (KZN): Distinguished by single row of suckers on arms (to date neither have been Size recorded on demersal surveys). Up to 360 mm mantle length and more than 10 kg. References Distribution Jereb et al., 2014; Nesis, 1987; Roper et al., 1984. West and South Coasts. Usually deeper than 100 m.

331 Phylum: Mollusca Cephalopoda

Octopus vulgaris (OctVul)

Phylum: Mollusca

Class: Cephalopoda

Order: Octopoda

Suborder: Incirrata

Family: Octopodidae

Common: Common octopus Alternate: Octopus “vulgaris” III Note: Octopus vulgaris is currently regarded as a single widely distributed species with a number of regional forms. These regional forms may be distinct species.

Lateral arms (Arms II & III) distinctly longer Skin “tight”, than medial arms (Arms I & IV) Ligula not forming a 2.5% arm fold at end of length mantle

Hectocotylised right Arm III

Colour usually with greyish tones Small, spoon-shaped ligula on hectocotylised right Arm III

100 mm

Distinguishing features Similar species • Ink sac present. Enteroctopus magniicus: All arms similar length, • Large muscular species; arms long, 4x to 5.5x lateral pairs (II & III) NOT distinctly longer than median ML; lateral pairs distinctly longer than median pairs; no enlarged suckers on lateral arms; generally pairs; Arm formula III ≥ II > IV > I. larger and found at greater depths; colour usually • Both sexes with two to three enlarged suckers with reddish rather than greyish tones; characteristic on lateral arms at level of 15th–19th proximal fold of loose skin at end of mantle; ligula prominent, suckers. long (16-25% of arm length), tapering to a blunt tip. • Colour usually with greyish tones. No loose skin Benthoctopus: Ventral surface of arms dark brown; fold at the end of the mantle. suckers prominent. Arm formula I = II = III = IV. • Four large erectile papillae in diamond arrangement on dorsal mantle. Bathypolypus: Small, purple with short arms; ink • One to two supraocular papillae over each eye. sac absent. Eledone schultzei (inshore, under 20 m depth) and Hectocotylus Velodona togata (KZN): Distinguished by single Right Arm III. Ligula small, spoon-shaped, 2.5% of row of suckers on arms (to date neither have been arm length. recorded on demersal surveys).

Size References Maximum weight 10 kg. Jereb et al., 2014; Nesis, 1987; Roper et al., 1984; Sanchez, 1988. Distribution West and South Coasts. To about 200 m, but generally less than 100 m.

332 Phylum: Mollusca Cephalopoda

Opisthoteuthis massyae (Opisto)

Phylum: Mollusca

Class: Cephalopoda

Order: Octopoda

Suborder: Cirrata

Family: Opisthoteuthidae

Common: Umbrella octopus Alternate: vossi

LATERAL VIEW ORAL VIEW

Paddle-like in Arms fully encased in Red eyes thick web In males: Dorsal arms (Arms I) more robust than Suckers in other arms single row

Suckers 5, 6 & 7 enlarged A row of cirri on either side of suckers

Arms fully encased in thick web 50 mm

Distinguishing features Distribution • Arms almost fully encased in a thick web with West and South Coasts between 500 and 1 500 m. a single row of suckers to the tips, lanked by a row of cirri on either side. A pair of small ins Similar species near posterior end of mantle. Four nominal species have been reported from • It looks like a dark reddish-brown gelatinous Namibia and South Africa: O. agassizi, O. grimaldii, blob, and it is only the eight rows of suckers on O. massyae and O. vossi. Villanueva et al. (2002) the oral side that show that it is a . revised the in the Atlantic, they designate • In males, the proximal four suckers on each arm O. vossi as a junior of O. massyae and restrict are small, the next three to six enlarged, then O. agassizi to the Caribbean and O. grimaldii to the decrease progressively to tips, but with a second eastern Atlantic from Azores to northern Namibia. ield of enlarged suckers at the web margin. This leaves O. massyae as the only known species • Dorsal arms (Arms I) of males thick, muscular of South Africa. Male O. grimaldii lack enlarged and robust to web margin, distal 3rd attenuate dorsal arms, females diicult to distinguish from and slender. Dorsal arms of females not O. massyae. O. grimaldii may be conined to deeper diferent to the other arms. water as all known specimens were collected between 1 135 and 2 287 m. Hectocotylus None. References Jereb et al., 2014; Sanchez, 1988; Sanchez & Guerra, Size 1989; Villanueva et al., 2002. 70 mm mantle length.

333 Phylum: Mollusca Cephalopoda

Vampyroteuthis infernalis (VamInf)

Phylum: Mollusca

Class: Cephalopoda

Order: Vampyromorpha

Suborder: -

Family: Vampyroteuthidae

Common: Alternate: -

Long thread-like ilament that can be retracted into pit on outer arm crown Stalked suckers Pair of ins near in single row posterior end of mantle

Cirri on each side of suckers Large pair of light organs at base of ins

Filamentous retractable limb Body black and gelatinous 10 mm

Illustration from Jereb et al., 2014 Badly damaged specimen captured in 2015

Distinguishing features Size • Eight arms, webbed for most of their length. Maximum 130 mm mantle length. • Two long ilamentous limbs that can be retracted into pits between Arms I and II. Distribution • A single row of stalked suckers on distal 2/3 of Mesopelagic (600-1 200 m) on West and arms, lanked by a row of cirri on either side. South Coasts. • Body gelatinous with black pigmentation. • A pair of small ins near posterior end of mantle Similar species in adults; juveniles with two pairs of ins. None, only known black octopod in the area.

Hectocotylus References None. Jereb et al., 2014; Young, 2009.

334 Phylum: Mollusca Cephalopoda

Spirula (Spirul)

Phylum: Mollusca

Class: Cephalopoda

Order: Spirulida

Suborder: -

Family: Spirulidae

Common: Ram’s horn squid Alternate: -

DORSAL VIEW VENTRAL VIEW

Mantle slightly Mantle deeply produced emarginate

Coiled internal shell

Small terminal Coiled internal shell ins 10 mm 5 mm

Distinguishing features Size • Tightly coiled, chambered shell. The shell is Maximum size 45 mm mantle length. internal, but visible on both dorsal and ventral surfaces. Distribution • Rectangular mantle with small ins at the Pelagic in surface waters on West and South Coasts. posterior corners. Seldom captured on demersal surveys, regular on • Mantle margin produced dorsally and deeply pelagic surveys. emarginated ventrally. • Colour dark reddish brown, but usually skinned Similar species during trawl capture. None.

Club References Small, marginally wider than the stalk. Suckers small, Jereb & Roper, 2010; Nesis, 1987. subequal.

Hectocotylus Both ventral arms modiied.

335 Phylum: Mollusca Cephalopoda

Quick guide to the Genus See Figure 6 (p. 325) for illustration of emarginated versus entire ventral mantle margin. If you are unsure of the species, but are sure that your Sepia is in the subgenus Hemisepius, then use the code “Hemisep”, otherwise use the code “Sepia”.

Table 1: Large Sepia – mainly South Coast Character Sepia papillata Sepia simoniana Sepia tuberculata Sepia vermiculata Dorsal margin broadly produced slightly produced broadly produced produced dorsally Ventral margin entire entire entire entire ( ) or emarginated ( ) Dorsal mantle rough, densely smooth, covered densely covered smooth; no covered with small with ine papillae with obvious tubercles or tubercles tubercles papillae Wrinkled patches present usually absent; present absent rarely present on mantle Diameter of equal to or greater no enlarged much less than much less than largest club than width of club suckers width of club width of club suckers

Table 2: Medium-sized Sepia (but beware of small individuals of above and of large Sepia faurei)

Character Sepia angulata Sepia australis Sepia hieronis Sepia hieronis Dorsal margin slightly produced produced dorsally strongly produced produced Ventral margin entire entire entire emarginated Dorsal mantle densely covered smooth smooth smooth with obvious tubercles Posterior spine absent large, obvious absent absent Enlarged club absent present absent absent suckers Arms I, II & III long, strongly not attenuated not attenuated not attenuated attenuated

Table 3: Subgenus Hemisepius: Small to medium-sized, characterised by the presence of a leshy ridge on sides of belly (visible as an iridescent blue line) and shell partially or completely chitinised Character Sepia dubia Sepia faurei Sepia robsoni Sepia sp. A Sepia cf. typica Dorsal margin straight straight or straight slightly convex slightly convex slightly convex Ventral margin deeply emarginated emarginated entire or entire or emarginated emarginated emarginated Dorsal mantle sparsely densely smooth smooth or smooth papillose, 2 covered with sparsely large wart-like papillae or papillose growths tubercles Ventral pores absent absent absent absent present Shell hard calciied hard centre, completely soft hard centre, hard centre, edges soft edges soft edges soft Dorsal arms suckers to tips tips devoid of distal ½ devoid suckers to tips suckers to tips suckers of suckers

336 Phylum: Mollusca Cephalopoda

Sepia angulata (SepAng)

Phylum: Mollusca

Class: Cephalopoda

Order: Sepiida

Suborder: -

Family: Sepiidae

Common: - Alternate: -

DORSAL VIEW VENTRAL VIEW Head and arms densely covered Arms I, II and III with large strongly attenuated tubercles No enlarged suckers

Dorsal mantle margin Ventral convex, not mantle produced margin entire

Club long, Dorsal mantle recurved with densely covered Narrow ins many small with large obvious 10 mm separated suckers tubercles posteriorly

Distinguishing features Size • Mantle short and broad. Dorsal margin convex, ML up to 100 mm ( ) and 120 mm ( ). not produced. Ventrally entire. • Fins narrow, rounded. Separate posteriorly. Distribution • Dorsal surface of mantle, head and arms densely Coastal to 350 m on South Coast, but all research covered with large, coarse papillae. survey records 100-110 m. • Ventral surface of mantle generally smooth, with widely scattered large papillae. Similar species • Arm suckers arranged in four series to tips. See Tables 1 and 2 (page 336). • Arms I to III long and strongly attenuated. The combination of strongly attenuated arms and Club dorsal surface densely covered with large papillae distinguishes this species from all except Sepia Long, slightly recurved, bearing numerous subequal tuberculata. Difers from S. tuberculata in longer, small suckers. thinner arms, absence of wrinkled patches on the Hectocotylus belly, and absence of enlarged suckers on the clubs. Not described. References None.

337 Phylum: Mollusca Cephalopoda

Sepia australis (SepAus)

Phylum: Mollusca

Class: Cephalopoda

Order: Sepiida

Suborder: -

Family: Sepiidae

Common: Southern cuttleish Alternate: -

DORSAL VIEW VENTRAL VIEW Small distal suckers

Dorsal margin of mantle Greatly produced enlarged Belly red when medial fresh, but fades suckers Ventral mantle to white after margin entire death Tentacular club

Robust Fin, widely posterior separated Robust spine posteriorly 10 mm posterior spine

Distinguishing features Distribution • Strong, robust spine on posterior end of Common on both West and South Coasts to 500 m, . but most abundant (90% of records) 60-200 m. • Mantle oval, dark purple dorsally. Ventral surface reddish-brown to orange when fresh, Similar species but on death fades to white with red centre. See Table 2 (page 336). • Mantle margin produced dorsally and straight Distinguished from other cuttleish in the region by (not emarginated) ventrally. reddish belly and robust posterior spine. • Suckers on arms in four rows. S. elegans: Recorded of Namibia is similar, has Club smaller spine and part of each arm (extent varies Short, somewhat recurved. Suckers arranged in between sexes) with suckers arranged in two rows. transverse rows, ive suckers per row; size varies markedly, smaller distally and four greatly enlarged References median suckers near proximal end. Augustyn et al., 1995; Jereb & Roper, 2005; Roeleveld, 1972; Sanchez, 1988. Hectocotylus Left ventral arm hectocotylised.

Size Up to 85 mm mantle length, 5 gram.

338 Phylum: Mollusca Cephalopoda

Sepia dubia (SepDub)

Phylum: Mollusca

Class: Cephalopoda

Order: Sepiida

Suborder: -

Family: Sepiidae

Common: - Alternate: Hemisepius dubia

DORSAL VIEW VENTRAL VIEW

Club small with Arms short, few small suckers suckers biserial, but they are widely spaced and appear uniserial

Ventral arms with very wide keel Margin deeply Margin emarginate straight Diagnostic pair of large wart-like growths

Shell hard, calciied Fleshy ridge 10 mm

Distinguishing features Distribution • A leshy ridge without pores on sides of ventral Very rare, known from only two specimens, mantle typical of the subgenus Hemisepius. 150-200 m. • Shell hard, well-calciied, unlike any others in the subgenus Hemisepius. Similar species • Mantle rounded, papillate, with two large See Table 3 (page 336). complex wart-like growths. Difers from all others in the subgenus Hemisepius • Mantle margin straight dorsally, deeply in possessing a hard, calciied shell, wide keels on emarginate ventrally. ventral arms and diagnostic skin growths. Additional • Wide keels on ventral arms. diferences are: • Arms with small suckers to the tips. Suckers biserial, but widely spaced so that they look S. faurei: Dorsal mantle densely covered with small as though they are uniserial. round papillae; tips of Arms I inger-like without suckers. Club S. robsoni: Shell reduced, soft without hard centre; Small, with few small subequal suckers. distal half of Arms I inger-like without suckers. Hectocotylus S. sp A.: Mantle broadly oval; dorsal margin convex; Not described. Only known specimens are female. ventral margin entire. S. cf. typica: 10-12 diagnostic obvious black pores Size ventrally. 17 mm mantle length. References Adam and Rees, 1966; Roeleveld, 1972. 339 Phylum: Mollusca Cephalopoda

Sepia faurei (SepFau)

Phylum: Mollusca

Class: Cephalopoda

Order: Sepiida

Suborder: -

Family: Sepiidae

Common: - Alternate: Hemisepius faurei

DORSAL VIEW VENTRAL VIEW Tips of Arms I inger-like, devoid Distal half of suckers of Arms I - III No enlarged attenuated suckers

Head densely covered Mantle margin with small straight or papillae slightly convex Margin deeply emarginate

Tentacular club without enlarged suckers

Dorsal mantle Narrow leshy Tentacular densely ridge without club covered with pores visible as small papillae thin iridescent 10 mm blue line

Distinguishing features Size • A leshy ridge on sides of ventral mantle typical Most small (20-30 mm ML), but specimens of over of the subgenus Hemisepius visible as a narrow 40 mm ML have been recorded. iridescent blue line without pores. • Shell thin, not calciied, but middle hard to the Distribution touch as in most Hemisepius. South Coast; from coast to 900 m. Rare and easily • Mantle broad, almost round. Dorsal margin overlooked. straight, deeply emarginate ventrally. • Dorsal surface of mantle, head and arms Similar species brownish, densely covered with small round See Table 3 (page 336). papillae. • Arm suckers small, globose and biserial. Arms Difers from others in the subgenus Hemisepius in I attenuated for distal half, tips inger-like, densely papillose dorsal mantle and from all other devoid of suckers. Sepia in the region except S. robsoni in having tips • Web between Arms I, II and III not reaching half of Arms I inger-like, devoid of suckers. of arm length. S. robsoni: Dorsal mantle and head smooth, or with few papillae around edges of shell. Shell thin, Club completely chitinous without hard central area. Broad and slightly recurved with 33 small suckers in transverse rows. Median suckers slightly larger References than lateral suckers. Roeleveld, 1972.

Hectocotylus Not described. 340 Phylum: Mollusca Cephalopoda

Sepia hieronis (SepHie)

Phylum: Mollusca

Class: Cephalopoda

Order: Sepiida

Suborder: -

Family: Sepiidae

Common: - Alternate: -

Mantle shape MALE DORSAL VIEW MALE VENTRAL VIEW No enlarged sexually dimorphic suckers

Dorsal arms Small smaller than tentacular ventral arms club without enlarged suckers

MALE: Mantle margin entire FEMALE: Margin MALE: Long dorsal emarginate mantle projection FEMALE: Shorter Pearl-white belly

Smoothly Tentacular rounded without club posterior spine 10 mm

Distinguishing features Size • Dorsal projection of mantle in males long, 80 mm mantle length. reaching to between eyes, shorter in females. • Ventral mantle margin entire in males, Distribution emarginated in females. West and South Coasts, between 40 and 550 m. • Fins a very narrow, inconspicuous fringe along mantle, separate posteriorly. Similar species • Dorsal arms shorter than ventral arms, with See Table 2 (page 336). suckers to the tips. • Suckers biserial on basal two-thirds of arms. This is the second commonest Sepia species after Proximal 3rd biserial (females) or quadriserial S. australis. (males). Sepia australis: Large, robust posterior spine. Fin • Dorsal colour reddish brown. Ventral colour wider. Belly red. Tentacular club larger with enlarged white, with reddish or orange border near base medial suckers. Suckers quadriserial for entire length of ins. No posterior spine. of all arms.

Club References Small, curved with ive to six transverse rows of Augustyn et al., 1995; Jereb & Roper, 2005; Roeleveld, numerous small subequal suckers. 1972; Sanchez, 1988.

Hectocotylus Left ventral arm. Modiied region about half of arm. Transversely wrinkled with minute lateral suckers.

341 Phylum: Mollusca Cephalopoda

Sepia papillata (SepPap)

Phylum: Mollusca

Class: Cephalopoda

Order: Sepiida

Suborder: -

Family: Sepiidae

Common: - Alternate: -

DORSAL VIEW

Arms not attenuated

Diameter VENTRAL VIEW of largest sucker equal to club width

Mantle Mantle margin broadly entire produced

Patches of wrinkled skin Skin rough, densely Tentacular tuberculate 50 mm club

Distinguishing features Size • Mantle broadly oval. Mantle margin dorsally 140 mm mantle length. produced, ventrally slightly emarginated. • Fins wide, rounded. Separate posteriorly. Distribution • Dorsal surface of mantle, head and arms densely Mainly South Coast shallower than 210 m. covered with small tubercles. • Wrinkled areas on ventral surface of mantle and Similar species on outer sides of ventral arms. See Table 1 (page 336). • Arm suckers not globose, arranged in four series basally, in four ( ) or eight ( ) rows on distal ¼. Wrinkled patches on belly unique to S. papillata and • Arms I to III not attenuated, webbed for about S. tuberculata and rarely S. simoniana. half of length. S. tuberculata: Dorsal surface of head and body • Colour: Dorsal dark reddish-brown to purple; densely covered with large, coarse tubercles. Club ventral pale with scattered . long, enlarged median suckers less than width of club. Arms I-III attenuated suckers in four rows to tip Club in both sexes. Maximum size 82 mm ML. Long. Small suckers distally. Four enlarged medial S. simoniana: Club very long, with numerous minute suckers. Middle two extremely large, diameter suckers. Normally lacks wrinkled patches on belly. approximately equal to width of the sucker-bearing surface of the club. References Hectocotylus Augustyn et al., 1995; Jereb & Roper, 2005; Roeleveld, 1972. Left ventral arm. In modiied region sucker rows two and three separated by naked area with transverse ridges.

342 Phylum: Mollusca Cephalopoda

Sepia robsoni (SepRob)

Phylum: Mollusca

Class: Cephalopoda

Order: Sepiida

Suborder: -

Family: Sepiidae

Common: - Alternate: Hemisepius robsoni

Arm tips without DORSAL VIEW VENTRAL VIEW suckers, inger-like Arms I, II & III Distal ½ of Arms I webbed half of inger-like, devoid length of suckers

Small, crescent- shaped club without enlarged suckers Ventral view of the dorsal arms Mantle margin Shell thin, lexible, slightly convex Mantle deeply completely emarginate chitinised

Wide ins along Fleshy ridge Dorsal mantle two-thirds of without smooth, with no mantle pores or few papillae

10 mm

Distinguishing features Size • A leshy ridge on sides of ventral mantle typical Maximum 20 mm. of the Subgenus Hemisepius visible as a narrow iridescent line without pores. Distribution • Shell thin, completely chitinous, lacking the Uncommon on both West and South Coasts, from hard centre of other Hemisepius. 300-500 m. • Mantle broad; dorsal margin convex, almost straight; ventral margin deeply emarginate. Similar species • Dorsal surface of mantle, head and arms brown, Other species in the subgenus Hemisepius covered with small, round papillae. (see Table 3 on page 336). • Arm suckers small, globose and biserial. Distal half of dorsal arms inger-like, devoid of suckers. S. dubia, S. sp. A and S. cf. typica: Shell with hard, Suckers to the tips of ventral and dorso-lateral calciied central area. Arms with suckers to the tips arms. and not attenuated. In addition, S. cf. typica ten to • Arms I, II & III webbed half of arm length. twelve pairs of obvious black pores ventrally. • Wide in not reaching edge of mantle (along S. faurei: Centre of shell hard; dorsal surface of mantle 60-80% of mantle) and separate posteriorly. densely covered with papillae or tubercles; distal half of Arms I-III attenuated and webbed for less than half Club of length; tips of Arms I devoid of suckers. Crescent-shaped, with about 53 subequal suckers in transverse rows of four to six. References Augustyn et al., 1995; Jereb & Roper, 2005; Roeleveld, Hectocotylus 1972; Sanchez, 1988. Left ventral arm. Ten pairs of minute suckers in modiied basal ¾.

343 Phylum: Mollusca Cephalopoda

Sepia simoniana (SepSim)

Phylum: Mollusca

Class: Cephalopoda

Order: Sepiida

Suborder: -

Family: Sepiidae

Common: - Alternate: -

DORSAL VIEW VENTRAL VIEW

Wrinkled patches

Mantle somewhat produced Margin entire

Club very long with minute suckers Usually Finely papillose smooth, rarely with Tentacular wrinkled club 50 mm patches

Distinguishing features Size • Mantle broadly oval. Mantle margin dorsally 185 mm mantle length. produced, ventrally entire. • Fins narrow, rounded. Separate posteriorly. Distribution • Skin inely papillose on dorsal surface of mantle, Mainly South Coast. Recorded to 190 m, but usually head and arms. less than 100 m. • Wrinkled areas on outer sides of ventral arms and rarely on ventral mantle. Similar species • All except ventral arms attenuated over distal See Table 1 (page 336). quarter. • Arm suckers not globose, quadriserial to tips of Long clubs with numerous small suckers are all arms in both sexes. diagnostic. • Colour: Dorsal pinkish-brown; ventral pale with S. vermiculata: Mantle broadly oval; slightly scattered chromatophores. produced. Skin dorsal and ventral smooth. Arms I-III attenuated tips. Club large, enlarged median Club suckers 3x of marginal. Very long, more than half the length of mantle, with S. papillata and S. tuberculata have large wrinkled numerous minute suckers. patches on the belly, and enlarged suckers on the clubs. Hectocotylus Left ventral arm. Modiied region – two ventral References and two dorsal rows of minute suckers separated Augustyn et al., 1995; Jereb & Roper, 2005; Roeleveld, by naked region with transverse ridges. Distal half 1972. normal.

344 Phylum: Mollusca Cephalopoda

Sepia sp. A (Sep001)

Phylum: Mollusca

Class: Cephalopoda

Order: Sepiida

Suborder: -

Family: Sepiidae

Common: - Alternate: Hemisepius sp. A

DORSAL VIEW VENTRAL VIEW

Numerous small Arms not Arm suckers subequal suckers attenuated small, and webbed biserial to for 50% of the tips length

Mantle margin Mantle convex margin entire or Tentacular Dorsal colour emarginate club tan with greenish and yellowish tones

Broad iridescent blue line Crescent- 10 mm without pores shaped club

Distinguishing features Distribution • A leshy ridge on sides of ventral mantle typical West and South Coasts, between 50 and 500 m. of the subgenus Hemisepius visible as a narrow iridescent blue line without pores. Similar species • Shell reduced. Middle hard to the touch, Other species in the subgenus Hemisepius margins soft, chitinous. (see Table 3 on page 336). For many years has • Mantle margin: slightly convex dorsally; been misidentiied as Sepia dubia. ventrally entire or shallowly emarginate. S. dubia: Very large keel on ventral arms; ventral • Arm suckers small and biserial. Dorsal arms not margin deeply emarginated; dorsal mantle sparsely attenuated and bearing suckers to the tips. papillate with two large wart-like growths on dorsal • Dorsal surface of mantle, head and arms mantle. greenish, with well-spaced round papillae. S. faurei: Dorsal mantle densely covered with small Club round papillae; tips of Arms I inger-like without Small, crescent-shaped, thicker than tentacle, with suckers. numerous small subequal suckers. S. robsoni: Shell reduced, soft without hard centre; distal half of Arms I inger-like without suckers. Hectocotylus Left ventral arm. S. cf. typica: 10-12 diagnostic obvious black pores ventrally. Size Up to 17 mm mantle length. References None.

345 Phylum: Mollusca Cephalopoda

Sepia tuberculata (SepTub)

Phylum: Mollusca

Class: Cephalopoda

Order: Sepiida

Suborder: -

Family: Sepiidae

Common: - Alternate: -

DORSAL VIEW VENTRAL VIEW

Arms I, II and III Largest sucker attenuated narrower than club width (see Sepia papillata) Head and arms densely covered with large tubercles

Dorsal mantle margin convex, Mantle not produced margin entire

Fin wide and rounded Patches of wrinkled skin on ventral Dorsal mantle mantle 10 mm densely covered with large, Fins separated Tentacular obvious tubercles 50 mm posteriorly club

Distinguishing features Size • Mantle short and broad. Dorsal margin convex, 82 mm mantle length. not produced. Ventrally entire. • Fins wide, rounded. Separate posteriorly. Distribution • Dorsal surface of mantle, head and arms densely Shallower than 200 m on South Coast. covered with large coarse tubercles. • Large wrinkled patches on either side of Similar species otherwise smooth ventral surface of mantle and See Table 1 (page 336). on outer area of ventral arms. Wrinkled patches on belly unique to S. papillata and • Arm suckers not globose, arranged in four series S. tuberculata and rarely S. simoniana. to tips. Tips of Arms I to III attenuated, webbed for less than half of arm length. S. papillata: Mantle produced dorsally; slightly emarginated ventrally; arms not attenuated, suckers Club in 4 ( ) or 8 ( ) rows on distal ¼; club large, diameter of enlarged median suckers equal to club width. Long, slightly recurved. Small suckers distally with enlarged suckers proximally. Diameter of largest S. simoniana: Club very long, with numerous minute suckers less than width of the sucker-bearing part suckers. of the club. S. angulata: Also has large, obvious tubercles on dorsal, but lacks wrinkled patches on belly. Hectocotylus Left ventral arm. The two dorsal rows of suckers References normal, separated from reduced ventral suckers by Augustyn et al., 1995; Jereb & Roper, 2005; Roeleveld, a broad naked area with transverse ridges. Distal 1972. half of arm normal.

346 Phylum: Mollusca Cephalopoda

Sepia cf. typica (SepTyp)

Phylum: Mollusca

Class: Cephalopoda

Order: Sepiida

Suborder: -

Family: Sepiidae

Common: - Alternate: Hemisepius cf. typica

DORSAL VIEW VENTRAL VIEW

Arms I with Small and suckers to the tip, straight. not attenuated Numerous small or inger-like subequal suckers

Margin convex Margin entire or emarginated

Mantle almost as wide as long Row of 10-12 pores in leshy ridge Tentacular diagnostic club Fin narrow, fused 10 mm posteriorly

Distinguishing features Hectocotylus • A leshy ridge on sides of ventral mantle typical Left ventral arm. Basal half modiied. Suckers minute, of the Subgenus Hemisepius, with 5-15 (usually widely spaced, separated by leshy transverse ridges. 10-12) diagnostic obvious black pores. • Shell not calciied, very thin and fragile, but hard Size to the touch. 25 mm mantle length. • Mantle very broadly oval, almost as wide as long; dorsal margin convex; ventral margin Distribution entire or emarginated. Both West and South Coasts, from coast to 600 m. • Dorsal surface of head and mantle greenish, sparsely papillose. Similar species • Suckers globose, biserial and extending to the See Table 3 (page 336). tips of the arms. Tips not attenuated. • Arms short, subequal in length; interbranchial Distinguished from all others in the Subgenus web between Arms I-III half arm length. Hemisepius (Sepia dubia, S. faurei, S. robsoni, and • Fins narrow, fused posteriorly. S. sp. A) by the presence of pores in the leshy ventral ridge. Club Small and straight, with numerous small subequal References suckers in transverse rows of six. Roeleveld, 1972.

347 Phylum: Mollusca Cephalopoda

Sepia vermiculata (SepVer)

Phylum: Mollusca

Class: Cephalopoda

Order: Sepiida

Suborder: -

Family: Sepiidae

Common: - Alternate: Sepia oicinalis vermiculata

DORSAL VIEW VENTRAL VIEW

Dorsal arms shortest

Ventral arms longest Mantle produced Margin emarginated (female) or entire (male)

No wrinkled patches on belly Skin smooth, without tubercles or papillae Fin wide and rounded

Small gap Tentacular separating ins club 100 mm

Distinguishing features Size • Mantle broadly oval. Dorsal margin convex, 287 mm mantle length. somewhat produced, ventral margin entire (male) or emarginated (female). Distribution • Fin wide, rounded. Along entire margin of Mainly shallow water on South Coast, but recorded mantle with small gap at tail. to 290 m. • Skin smooth, both dorsally and ventrally, no obvious pores or wrinkled patches. Similar species • Ventral arms longest, dorsal arms shortest. Arms See Table 1 (page 336). III and IV keeled, joined by shallow web. • Suckers on arms in four rows, extending to Adults can be confused only with the other somewhat attenuated tips. three large species, S. papillata, S. simoniana and • Some individuals show diagnostic transverse S. tuberculata. zebra-like stripes on mantle and ventral arms. S. simoniana: Dorsally mantle more produced and skin finely papillose. Arms I-III attenuated and Club webbed for about half of length. Club very long, Large, one third of mantle length; distal suckers with numerous minute suckers. small in oblique rows of eight; proximal suckers in S. papillata and S. tuberculata have large wrinkled oblique rows of ive, with median suckers 1.5-2 times patches on the belly. and middle suckers 3 times the size of the marginal suckers. References Hectocotylus Jereb & Roper, 2005; Roeleveld, 1972; Sanchez, 1988. Left ventral arm. Modiied region with 9-12 rows of reduced suckers separated by transverse ridges. 348 Phylum: Mollusca Cephalopoda

Austrorossia enigmatica (RosEni)

Phylum: Mollusca

Class: Cephalopoda

Order: Sepiida

Suborder: -

Family: Sepiolidae

Common: Alternate: enigmatica; Austrorossia mastigophora

DORSAL VIEW VENTRAL VIEW Numerous small subequal suckers

Tentacular club long Very large Constriction and narrow, green eyes in front of with small eyes suckers Mantle not Tentacular Shell extended as a club rudimentary Large, ventral shield rounded ins attached midway along the mantle

10 mm Iridescent green eye

Distinguishing features Size • Shell rudimentary, chitinous, feels as though 40 mm mantle length. there is no internal shell. • Fins large, rounded, attached about midway Distribution along mantle; broadly separated posteriorly. West (common) and South (uncommon) Coasts, • Anterior mantle edge not fused with head between 200 and 500 m. dorsally, not covering funnel ventrally. • Head short and broad, constricted round crown Similar species of circumoral appendages anterior to eyes. None. • Eyes large, prominent iridescent green. References Club Nesis, 1987; Sanchez, 1988. Narrow, not wider than tentacle. Suckers microscopic in 30-40 rows.

Hectocotylus Both dorsal arms.

349 Phylum: Mollusca Cephalopoda

Inioteuthis capensis (Inio)

Phylum: Mollusca

Class: Cephalopoda

Order: Sepiida

Suborder: -

Family: Sepiolidae

Common: - Alternate: Rondeletiola capensis

DORSAL VIEW VENTRAL VIEW

Dorsal mantle fused to head by a narrow band Anterio- ventral edge Small rounded of mantle ins attached far not extended forward as a ventral Mantle cut shield open to show absence of ink sac Fins very widely photophores Mantle separated oblong posteriorly

10 mm

Distinguishing features Size • Shell absent. 20 mm mantle length. • Fins small, rounded, attached mid-laterally to mantle. Broadly separated posteriorly. Distribution • Mantle fused with head dorsally by a narrow Common in surface waters on both coasts, but occipital band. seldom recorded on demersal surveys because of • Funnel not covered by a forward extension of small size. the anterio-ventral edge of mantle. • Body oblong, longer, less eyeball-like than Similar species Stoloteuthis. Stoloteuthis: Larger, more globular; dorsal mantle • Ventral surface of ink sac without luminous broadly fused with head (occipital band 40-50% organ. head width); ventral mantle extended to form a ventral shield; luminous organ present on ink sac. Club Small, slightly wider than stalk, with small suckers. References Nesis, 1987. Hectocotylus Left dorsal arm. Basal part modiied into specialised copulatory apparatus.

350 Phylum: Mollusca Cephalopoda

Stoloteuthis sp. (Stolot)

Phylum: Mollusca

Class: Cephalopoda

Order: Sepiida

Suborder: -

Family: Sepiolidae

Common: Eye-ball squid, Butterly bobtail squid Alternate: -

DORSAL VIEW VENTRAL VIEW

Mantle extended to eye level Top of head as a ventral iridescent shield

Ventral view with Dorsal mantle folded back to expose the mantle Round, on the fused to globular ink sac head for body 40-50% Light head width organ on ink sac

Small dorso- lateral ins attached far back 10 mm

Distinguishing features Hectocotylus • Shell absent. Both Arms II. • Fins large, ear-like, attached laterally to posterior half of mantle. Broadly separated posteriorly. Size • Dorsal mantle edge fused to head by a broad 20 mm mantle length. occipital band 40-50% of head width. • Anterior edge of mantle extended as a ventral Distribution shield to level with eyes. Between 100 and 850 m, mainly on West Coast. • Body round, globular, looks like an eyeball. Top of head iridescent green. Similar species • First three pairs of arms joined by a deep web. Inioteuthis capensis: Smaller, body longer, less Suckers on arms in two series. globular. Anterior edge of mantle not extended • A luminous organ on ventral side of ink sac. into a ventral shield. Mantle narrowly fused to head dorsally. No luminous organ on the ink sac. Club Not thicker than tentacle, with numerous small References suckers. Nesis, 1987.

351 Phylum: Mollusca Cephalopoda

Afrololigo mercatoris (Lollig)

Phylum: Mollusca

Class: Cephalopoda

Order: Myopsida

Suborder: -

Family:

Common: African thumbstall squid Alternate: mercatoris

DORSAL VIEW Reddish spots on mantle, head and arms

Dorsal arms short

Fins small, posterior margin Ventral mantle deeply 5 mm convex emarginated, edge extended in lateral points

Tentacular club

VENTRAL VIEW

Distinguishing features Size • Lens of eye covered by a cornea, not in direct Males 50 mm mantle length, females 35 mm. contact with seawater. • Funnel-locking cartilage a simple straight Distribution groove and ridge. West and South Coasts to 470 m. • Arms with two rows of suckers, clubs with four rows. No hooks. Similar species • Dorsal arms (Arms I) much shorter than other Juvenile reynaudii have longer, narrower ins arms. with concave posterior margins; ventral mantle • Fins translucent, short (40% ML) and rounded, shallowly emarginated; and lack the irregular reddish with convex posterior margins. spots. Clubs wider, with some enlarged suckers. • White, with irregular reddish-brown spots on mantle, head and arms. References Jereb & Roper, 2010; Nesis, 1987; Sanchez, 1988. Club Narrow, small, with suckers arranged in four longitudinal rows; four to ive pairs of medial suckers on manus enlarged, sucker rings with 15-25 teeth.

Hectocotylus Left ventral arm. Basal half normal; distal half with elongate papillae.

352 Phylum: Mollusca Cephalopoda

Loligo reynaudii (CHOK)

Phylum: Mollusca

Class: Cephalopoda

Order: Myopsida

Suborder: -

Family: Loliginidae

Common: Chokka Alternate: Loligo vulgaris reynaudii

VENTRAL VIEW Suckers on manus greatly expanded Fin rhomboidal with concave posterior Mantle irm margin

Suckers on arms biserial

Skin red, deciduous, Fin long, more Eye covered by a usually rubbed of than 65% of cornea Suckers in four rows, in the trawl showing mantle those on manus white mantle greatly expanded; cups smooth without teeth Damaged specimen 50 mm

Tentacular club large, without hooks

Distinguishing features Size • Lens of eye covered by a cornea, not in direct Males up to 400 mm mantle length. Females smaller. contact with seawater. • Funnel-locking cartilage a simple straight Distribution groove and ridge. Widespread on both coasts but most common on • Arms with two rows of suckers, clubs with four South Coast, shallower than 200 m. rows. No hooks. • Mantle narrow, irm and elongate. Similar species • Skin red, deciduous, usually rubbed of in the Juveniles can be confused with Afrololigo mercatoris trawl. of similar size. • Fins posterior, long, over 65% of mantle, rhomboidal in shape, with concave posterior Afrololigo mercatoris: Short, rounded fins with margin. convex posterior margin; ventral mantle deeply emarginated; clubs small, narrow, without greatly Club enlarged suckers on manus; mantle and arms with reddish spots. Tentacles long; clubs expanded; suckers in four series; suckers on manus greatly enlarged, cups duvaucelii from KwaZulu-Natal which has smooth without chitinous teeth. a wider club with four rows of enlarged suckers.

Hectocotylus References Left ventral arm. Basal part of arm with two series Jereb & Roper, 2010; Nesis, 1987; Sanchez, 1988. of suckers. Suckers on distal part reduced, but with elongated stalks to form papillae making a feathery tip.

353 Phylum: Mollusca Cephalopoda

Chtenopteryx sicula (CteSic)

Phylum: Mollusca

Class: Cephalopoda

Order: [Unassigned]

Suborder: -

Family: Chtenopterygidae

Common: Comb-inned squid Alternate: Ctenopteryx sicula (common misspelling)

DORSAL VIEW Photogenic patches on ventral surface of eyeballs Fins with muscular ribs creating a comb-like appearance

Arms I to III with small suckers in transverse series of 6 to 14 Clubs slender with minute suckers Figure reproduced from Jereb & Roper, 2010, with permission.

Distinguishing features Club • Eye not covered by a transparent membrane, Narrow, not expanded, with minute suckers in 8 to lens in open contact with seawater. 20 irregular transverse series. • Funnel-locking cartilage a simple straight groove and ridge. Hectocotylus • Mantle broadly rounded with ins along full None. length, similar shape to cuttleish. • Fins comprised of slender rib-like muscle Size bundles connected by membranes, giving a Up to 100 mm mantle length. comb-like appearance. • Arms I to III with suckers in transverse series of Distribution 6 to 14; Arms IV with a few small suckers in a Both West and South Coasts, from 500 to 1000 m. zigzag pattern. • Large photogenic patches on ventral surface of Similar species eyeballs. Comb-like ins are diagnostic. • Minute suckers on lappets of the buccal membrane. References Jereb & Roper, 2010.

354 Phylum: Mollusca Cephalopoda

Ancistrocheirus lesueurii (AncLes)

Phylum: Mollusca

Class: Cephalopoda

Order: Oegopsida

Suborder: -

Family: Ancistrocheiridae

Common: Sharpear enope squid Alternate: Thelidioteuthis alessandrinii

DORSAL VIEW Note: Usually badly damaged in trawl, completely VENTRAL VIEW skinned and the head usually separated from the body.

Mantle elongate forming a tail

Hooks present Fin very large, on all arms subterminal with characteristic brown transverse dashed lines Illustration from Jereb & Roper (2010) showing 20-24 large 100 mm scattered photophores

Distinguishing features Hectocotylus • Eye not covered by a transparent membrane, Right ventral arm. lens in open contact with seawater. • Funnel-locking cartilage a simple straight Size groove and ridge. Attains 410 mm mantle length, and 3 kg. • Fin very large, rhomboidal, length 70-80% and width 80% of ML. Attached from anterior of Distribution mantle ending slightly subterminal. Dorsal Mesopelagic and bathypelagic on West and South surface with irregular brownish transverse Coasts. dashed lines (photophores). • Posterior end of mantle elongated, forming a Similar species tail. also have very large rhomboidal • Arms robust with two series of hooks. Small ins, but lack the brown dashed lines on the dorsal suckers sometimes present on tips. surface, and the tentacles are reduced or absent. • Ventral surface of mantle studded with 20-24 relatively large separated photophores. No References photophores on eyeballs or viscera. Jereb & Roper, 2010; Nesis, 1987; Sanchez, 1988. Club Tentacles robust, 12 photophores on aboral side of stalk. Clubs not expanded, two series of hooks on manus, no suckers. Discrete carpal cluster.

355 Phylum: Mollusca Cephalopoda

Brachioteuthis picta (BraPic)

Phylum: Mollusca

Class: Cephalopoda

Order: Oegopsida

Suborder: -

Family:

Common: Ornate arm squid Alternate: -

DORSAL VIEW

Abrupt narrowing Sausage-shaped photophore of mantle on ventral surface of eye

Skin smooth Flared mantle Large in, length and opening width ± 50% ML Tentacular club Figure reproduced from Jereb & Roper, 2010, with permission.

Distinguishing features Hectocotylus • Eye not covered by a transparent membrane, Not described. lens in open contact with seawater. • Funnel-locking cartilage a simple straight Size groove and ridge. 90 mm mantle length. • Mantle long and slender; slightly lared at opening; abruptly narrows anterior to ins. Distribution • Fin almost as wide as long, length and width Oceanic on both West and South Coasts. about 50% ML. • A sausage-shaped photophore on ventral Similar species surface of each eye. sp. A. has rough “warty” skin. Fin • Skin smooth in both sexes, never rough even in length less than 50% ML, width greater than length. mature individuals. References Club Jereb & Roper, 2010; Lipinski, 2001; Nesis, 1987; Manus greatly expanded, covered with numerous Sanchez, 1988. rows of small, long-stalked suckers; dactylus section with three to four rows of suckers.

356 Phylum: Mollusca Cephalopoda

Brachioteuthis sp. A (Brachi)

Phylum: Mollusca

Class: Cephalopoda

Order: Oegopsida

Suborder: -

Family: Brachioteuthidae

Common: - Alternate: -

DORSAL VIEW

Large clubs Dactylus without hooks

Flared mantle Abrupt narrowing Small opening of mantle suckers without hooks Manus

Sausage-shaped photophore Granular skin on Tentacular club showing ventral surface of eye greatly expanded manus and elongated dactylus

50 mm

Distinguishing features Hectocotylus • Eye not covered by a transparent membrane, Not described. lens in open contact with seawater. • Funnel-locking cartilage a simple straight Size groove and ridge. 90 mm mantle length. • Mantle long and slender; slightly lared at opening; abruptly narrows anterior to ins. Distribution • Fin length less than 50% of ML. Width greater Mesopelagic on both West and South Coasts, deeper than length. than 300 m. • A sausage-shaped photophore on ventral surface of each eye. Similar species • Skin rough, granular. : Smooth skin; in length equals width, about 50% of ML. Club Manus greatly expanded, covered with numerous species also have rough, warty skin but difer rows of small, long-stalked suckers. Dactylus section in the presence of hooks on the clubs. with three to four rows of suckers. References Jereb & Roper, 2010; Lipinski, 2001; Nesis, 1987.

357 Phylum: Mollusca Cephalopoda

Chiroteuthis mega (ChrCap)

Phylum: Mollusca

Class: Cephalopoda

Order: Teuthoidea

Suborder: Oegopsida

Family:

Common: Atlantic long-arm squid Alternate: capensis

LATERAL VIEW Terminal light organ

Club Circular in fringed by Both head and lappets neck very long

Club of C. veranii (Jereb & Short “tail” Ink sac Roper, 2010) bearing a single Tentacle very long photophore and thin One One post- Ventral arms photophore ocular much larger on anterior photophore than others eye margin 50 mm Seven photophores on ventral eye margin

Distinguishing features Hectocotylus • Eye not covered by a transparent membrane, Absent. lens in open contact with seawater. • Funnel-locking cartilage oval, with two knobs Size directed towards centre of the concavity. 100-200 mm mantle length. • Head and neck long and tubular, but squarish near the eyes; head plus neck almost as long as Distribution mantle; eyes about midway between arm bases West and South Coasts, from 700 to 1400 m. and mantle. • Fins thick and leshy, together circular, not Similar species lobed; extends as a short tail past ins. Chiroteuthis veranii (possible occurrence on South • Arms IV much longer and thicker than other Coast) difers in having two photophores on the arms; Arms II and III subequal; Arms I short. ink sac. • Eyeball with one photophore on anterior margin just above midline and one just below : Long slender tail (greater than midline on posterior margin; a series of seven ML) posterior to short round in. Arms I - III very long, photophores on anterio-ventral margin. 2x mantle length and 3x length of Arms IV. Lacks • A single photophore on the ink sac. photophores on eyeballs.

Club References Tentacles long and very thin; club fringed with Jereb & Roper, 2010; Nesis, 1987; Sanchez, 1988; lappets; suckers in four rows arranged in pairs at Voss, 1967. the base of each lappet; a long oval light organ at the tip of the club.

358 Phylum: Mollusca Cephalopoda

Cranchia scabra (CrnScb)

Phylum: Mollusca

Class: Cephalopoda

Order: Oegopsida

Suborder: -

Family:

Common: Rough cranch squid Alternate: -

VENTRAL VIEW Ventral mantle with two cartilaginous strips extending from apex of each mantle-funnel fusion

Funnel fused to Fins nearly head laterally oval; posterior lobes free One between the inner and outer groups Mature females have a photophore at end of each arm Six around Seven in pupil arc on 14 small round ventral photophores on eyes Mantle sac-like Mantle fused to head dorsally edge with cartilaginous and to funnel ventrally tubercles Ventral view of eye showing arrangement of the 14 photophores 10 mm

Distinguishing features Hectocotylus • Eye not covered by a transparent membrane, Right ventral arm. Suckers in four series on midpoint lens in open contact with seawater. of hectocotylised arm. • Mantle fused to posterior-lateral corners of funnel, no funnel-locking cartilage. Size • Head fused to the mantle at the nuchal Up to 150 mm mantle length. cartilage. • Ventral surface of mantle with two cartilaginous Distribution strips extending posteriorly from anterior apex Both West and South Coasts, from 400 to 1200 m. of each funnel-mantle fusion; funnel fused to head laterally. Similar species • Brachial photophore on end of each arm in Monotypic genus. Cartilaginous tubercles scattered mature females. over entire mantle unique among Cranchiidae. • Eyes with 14 small round photophores: an inner similar in general shape, group of six around pupil; an outer group of but with smooth skin and funnel free from head seven in an arc on ventral edge; one between laterally. the inner and outer groups. • Mantle a thin-walled sac covered in spiky, References cartilaginous tubercles. Jereb & Roper, 2010; Nesis, 1987; Sanchez, 1988. • Fins small (less than 25% ML) posterior. Each nearly oval with free posterior lobe.

Club Not expanded, with small sub-equal suckers. Alternating series of carpal suckers and pads for most of tentacle length. 359 Phylum: Mollusca Cephalopoda

Leachia cyclura (LeaCyc)

Phylum: Mollusca

Class: Cephalopoda

Order: Oegopsida

Suborder: -

Family: Cranchiidae

Common: Leach’s cranch squid Alternate: -

DORSAL VIEW One cartilaginous strip extends ventrally for 20-30% of ML from apex of each funnel-mantle fusion

Cartilaginous dorsal keel with small tubercles along dorsal midline Mantle fused to head dorsally and to funnel ventrally

Mantle spindle- shaped, tapering to Eight small oval point Fins terminal, photophores on eyes rounded and fused posteriorly 10 mm

Distinguishing features Hectocotylus • Eye not covered by a transparent membrane, Not described. lens in open contact with seawater. • Mantle fused to posterior-lateral corners of Size funnel, no funnel-locking cartilage. Maximum 150-200 mm mantle length. • Head fused to the mantle at the nuchal cartilage. Funnel fused to head laterally. Distribution • Mantle spindle-shaped, tapering to sharp point, South Coast to west of Cape Point, from surface to with a cartilaginous dorsal keel. 2 000 m. • Ventral surface of mantle with one cartilaginous strip extending posteriorly for 20-30% of mantle Similar species length from anterior apex of each funnel-mantle atlantica: Cartilaginous strip 14-15% of ML; fusion. six photophores on each eye (ive outer and one • Body translucent with scattered chromatophores near pupil). and three dark internal organs easily visible. • Large elongate brachial photophore on tips of sp.: Head nearly as wide as mantle; either Arms III in mature females. 4 or 14 oval photophores on eyes depending on • Eight eye photophores, ive in outer row and species. three near pupil. • Fins terminal, rounded and fused posteriorly. References Jereb & Roper, 2010; Nesis, 1987. Club Median suckers on manus greatly enlarged.

360 Phylum: Mollusca Cephalopoda

Liocranchia reinhardti (LioRei)

Phylum: Mollusca

Class: Cephalopoda

Order: Oegopsida

Suborder: -

Family: Cranchiidae

Common: Reinhart’s cranch squid Alternate: -

VENTRAL VIEW If unsure of the species, use the code Liocra for Liocranchia sp.

Pair of moderately long Cartilaginous tubercles Mantle spindle- cartilaginous strips, studded along dorsal midline shaped, tapering to with tubercles extending in point V-shape from apex of lateral funnel-mantle fusion

Photophores at tips of Arms III in mature females

Mantle smooth

14 small photophores Mantle fused to head Fins terminal, rounded on eyeballs, arranged dorsally and to funnel and fused posteriorly in 3 groups ventrally

Ventral view (Vos, 1980) reproduced with the permission of the Bulletin of Marine Science

Distinguishing features Club • Eye not covered by a transparent membrane, Slightly expanded with small, sub-equal suckers. An lens in open contact with seawater. alternating series of carpal suckers and pads for most • Mantle fused to posterior-lateral corners of of tentacle length. funnel, no funnel-locking cartilage. • Mantle spindle-shaped, tapering to sharp point; Size cartilaginous tubercles along dorsal midline. Maximum 250 mm mantle length. • Head fused to the mantle at the nuchal cartilage. Funnel fused to head laterally. Distribution • Ventral surface of mantle with a pair of Pelagic to mesopelagic on West and South Coasts. cartilaginous strips, studded with tubercles, extending posteriorly in a V-shape from apex of Similar species each lateral funnel-mantle fusion (four strips in Liocranchia valdiviae: No cartilaginous tubercles on total). dorsal midline, four small round photophores on • Mature females with brachial photophores on eyes. tips of Arms III only. • 14 small photophores around eyes: four Leachia sp.: Head small, much narrower than width around pupil; eight in ventral arc; two between of mantle; one cartilaginous strip from each lateral the two series. funnel-mantle fusion; six or eight oval photophores • Fins terminal, rounded and fused posteriorly. on eyes depending on species.

Hectocotylus References Right or left ventral arm. Suckers in two series on Jereb & Roper, 2010; Nesis, 1987; Vos, 1980. midpoint of hectocotylised arms.

361 Phylum: Mollusca Cephalopoda

Liocranchia valdiviae (LioVal)

Phylum: Mollusca

Class: Cephalopoda

Order: Oegopsida

Suborder: -

Family: Cranchiidae

Common: Valdivia cranch squid Alternate: -

DORSAL VIEW Ventral mantle with two pairs of moderately long cartilaginous strips studded with tubercles, one pair extending in V-shape from apex of each lateral funnel-mantle fusion. If unsure of the species, use the code Liocra for Liocranchia sp.

No cartilaginous tubercles Mantle Mantle fused to head dorsally on dorsal mantle surface smooth and to funnel ventrally

No illustration Mature available, igure females have of L. reinhardti a photophore used to indicate at end of features. Figure Arms III only reproduced from Jereb & Roper, 2010, with permission. Fins terminal, Mantle spindle- Four small oval photophores rounded and fused shaped, tapering on each eye posteriorly to point

Distinguishing features Hectocotylus • Eye not covered by a transparent membrane, Right or left ventral arm. Suckers in two series on lens in open contact with seawater. midpoint of hectocotylised arms. • Mantle fused to posterior-lateral corners of funnel, no funnel-locking cartilage. Size • Head fused to the mantle at the nuchal Maximum 250 mm mantle length. cartilage. Funnel fused to head laterally. • Mantle spindle-shaped, tapering to sharp point, Distribution without cartilaginous tubercles along dorsal Pelagic to mesopelagic on West and South Coasts. midline. • Ventral surface of mantle with a pair of Similar species cartilaginous strips, studded with tubercles : Cartilaginous tubercles along extending posteriorly in a V-shape from apex of dorsal midline. Fourteen oval photophores on eye. each lateral funnel-mantle fusion (four strips in total). Leachia sp.: Head small, much narrower than width • Brachial photophore only on Arms III of of mantle; one cartilaginous strip from each lateral mature females. Eyes with four small round funnel-mantle fusion, six or eight oval photophores photophores. on eyes depending on species. • Fins terminal, rounded and fused posteriorly. References Club Jereb & Roper, 2010; Nesis, 1987. Slightly expanded with small, sub-equal suckers. An alternating series of carpal suckers and pads for most of tentacle length.

362 Phylum: Mollusca Cephalopoda

Megalocranchia maxima (Megalo)

Phylum: Mollusca

Class: Cephalopoda

Order: Oegopsida

Suborder: -

Family: Cranchiidae

Common: Large cranch squid Alternate: -

VENTRAL VIEW

Mantle long, slender, Mantle fused to head dorsally Adult MALES: tapering to point and to funnel ventrally Arm tips I-III Fin does not abruptly extend past attenuated gladius with minute suckers

Adult FEMALES: Arm tips I-III (or only Arm III) with Anterior 10-15% of Large complex, bilobed Lower half of eye covered by two photophore in fused to mantle photophore on ink sac photophores, one large crescent- shaped, the other small

Figure reproduced from Jereb & Roper, 2010, with permission.

Distinguishing features Hectocotylus • Eye not covered by a transparent membrane, Absent. lens in open contact with seawater. • Mantle fused to posterior-lateral corners of Size funnel, no funnel-locking cartilage. 1 800 mm mantle length. • Head fused to the mantle at the nuchal cartilage; funnel free from the head laterally. Distribution • Mantle slender, elongate, tapers dramatically to West and South Coasts, 600-2000 m during day; thin sharp tip, lacking cartilaginous tubercles. migrates to 100-700 m at night. • Most of ventral hemisphere of eyes covered by two photophores, a large crescent-shaped Similar species posterior photophore and a smaller anterior Distinguished from other Cranchiids except photophore within its concavity. by fusing of anterior of in to lateral • Large, complex, bilobed compound photophore mantle. present on ventral surface of rounded digestive gland and ink sac. Teuthowenia: Lack photophore on ink sac, have • Long lanceolate ins (50% ML) terminal-lateral three photophores on eyes and in extends beyond without anterior lobes; anterior 10-15% of in gladius. fused to lateral margins of mantle (unique to and Teuthowenia). References Jereb & Roper, 2010; Nesis, 1987. Club Carpal suckers in two series on tentacular stalk; clubs moderate, slightly expanded with suckers in four series. 363 Phylum: Mollusca Cephalopoda

Sandalops melancholicus (SanMel)

Phylum: Mollusca

Class: Cephalopoda

Order: Oegopsida

Suborder: -

Family: Cranchiidae

Common: Melancholy cranch squid Alternate: -

DORSAL VIEW Mantle fused to head Eye photophores: dorsally and to funnel one small anterior and ventrally one large posterior Mantle smooth, no cartilaginous tubercles

No brachial Fins small, photophores on subterminal arm tips

Head small, eyes large and bulbous Figure reproduced from Jereb & Clubs without hooks or Roper, 2010, with permission. enlarged suckers

Distinguishing features Size • Eye not covered by a transparent membrane, Maximum mantle length 110 mm. lens in open contact with seawater. • Mantle fused to posterior-lateral corners of Distribution funnel, no funnel-locking cartilage. Mesopelagic and bathypelagic on West Coast. • Head fused to the mantle at the nuchal cartilage; funnel free from the head laterally. Similar species • Mantle without cartilaginous tubercles, skin is superficially similar, but that smooth. species has rough skin, and funnel fused to head • Head small; eyes large, bulbous with two laterally. photophores (one large posterior and one small anterior). References • Arms with biserial, spherical suckers. Jereb & Roper, 2010; Nesis, 1987. • Fins small (12-15% ML), rounded, subterminal.

Club Club moderate, without enlarged suckers or hooks. Suckers in four series.

Hectocotylus Absent.

364 Phylum: Mollusca Cephalopoda

Taonius pavo (Taonis)

Phylum: Mollusca

Class: Cephalopoda

Order: Oegopsida

Suborder: -

Family: Cranchiidae

Common: Peacock cranch squid Alternate: -

DORSAL VIEW Mantle fused to head dorsally and to funnel ventrally

Mantle and in extended into Mantle very long and thin tail slender without tubercles

Fin long, narrow, strongly Clubs without Head small, eyes attenuated hooks large, bulbous with two large photophores

Enlarged suckers with one 50 mm or two hook-like teeth

Distinguishing features Hectocotylus • Eye not covered by a transparent membrane, Absent. lens in open contact with seawater. • Mantle fused to posterior-lateral corners of Size funnel, no funnel-locking cartilage. 650 mm mantle length. • Head fused to the mantle at the nuchal cartilage. Distribution • Funnel free from the head laterally. West and South Coasts. Juveniles below 600 m, • Mantle without cartilaginous tubercles, very adults to 2000 m. long, slender, tapering to long thin “tail”. • Head small; eyes large, bulbous with one large Similar species posterior crescent-shaped photophore that None. engulfs the small anterior photophore. • Arms with biserial, spherical suckers; without References hooks. Jereb & Roper, 2010; Nesis, 1987; Sanchez, 1988. • Fins long (50% ML), narrow, lanceolate, very attenuated posteriorly. Anterior lobes small.

Club Moderate without hooks; enlarged suckers with one or two large hook-like teeth.

365 Phylum: Mollusca Cephalopoda

Teuthowenia pellucida (Teuthw)

Phylum: Mollusca

Class: Cephalopoda

Order: Oegopsida

Suborder: -

Family: Cranchiidae

Common: - Alternate: -

DORSAL VIEW

Mantle fused to head dorsally Fin extends and to funnel ventrally past gladius as small lobes FEMALE: Light organs on tips of Arms I-IV

Carpal suckers Anterior 10-15% of Large bulbous eyes with Short, muscular in four series in in fused to mantle three photophores tentacles zigzag pattern

Figure reproduced from Jereb & Roper, 2010, with permission.

Distinguishing features Hectocotylus • Eye not covered by a transparent membrane, Absent. lens in open contact with seawater. • Mantle fused to posterior-lateral corners of Size funnel, no funnel-locking cartilage. 210 mm mantle length. • Head fused to the mantle at the nuchal cartilage. Distribution • Mantle thin, leathery; funnel free from the head West and South Coasts. Occur at greater depths laterally. with age; juveniles and subadults to 1 000 m; adults • Funnel/mantle fusion cartilages small, oval with 1 000-2 500 m. one to four cartilaginous tubercles at mantle margin. Similar species • Head small; eyes large, bulbous, with three Distinguished from other Cranchiids except nested photophores – a large crescent-shaped Teuthowenia by fusing of anterior of in to lateral posterior photophore, within its concavity a mantle. smaller crescent-shaped anterior photophore and a third small oval photophore. Megalocranchia: Has complex photophore on ink • Brachial end-organ (photophore) on tips of sac, and two on eyes; in does not extend beyond Arms I-IV of mature females. gladius. • Fins long, narrow, terminal-lateral, taper posteriorly, terminating in small lobes that References extend posteriorly beyond the tip. Jereb & Roper, 2010; Nesis, 1987.

Club Tentacles short, muscular; carpal suckers in four series in a zigzag pattern on stalk; club slightly expanded with suckers on long pedestals. 366 Phylum: Mollusca Cephalopoda

Discoteuthis discus (DisDis)

Phylum: Mollusca

Class: Cephalopoda

Order: Oegopsida

Suborder: -

Family:

Common: Discus squid Alternate: -

DORSAL VIEW

Tentacles well-developed Tentacular club Armature on (from Jereb & arms and clubs Roper, 2010) suckers only, no hooks

Funnel-locking cartilage

Suckers globose, biserial Fin large, disc-like, width greater than length Figure reproduced from Jereb & Roper, 2010, with permission.

Distinguishing features Distribution • Eye not covered by a transparent membrane, Rare. Possible on both West and South Coasts, lens in open contact with seawater. 500 to 1000 m. • Fin large, disc-like, wider than long, equal to mantle length. Similar species • Suckers on arms globose, biserial. Combination of large disc-like fin and globose • A single photophore on ventral mantle near suckers unique in area. Other species with large posterior end of body; no photophores on head in are: or near anterior edge of mantle, or on ink sac. Mastigopsis hjorti: Has small suckers, weak tentacles • Funnel-locking cartilage triangular, with an and two photophores on eyeball. oblique groove. Octopoteuthidae ( sicula and Club danae): Tentacle residual or absent; armature of Compact, widened, with four rows of suckers, two hooks. central rows greatly enlarged, globose. leseuerii: Armature of hooks, diagnostic dashed brown line on ins. Hectocotylus Absent. References Jereb & Roper, 2010; Nesis, 1987; Sanchez, 1988. Size 600 mm mantle length.

367 Phylum: Mollusca Cephalopoda

Abraliopsis gilchristi (AbrGil)

Phylum: Mollusca

Class: Cephalopoda

Order: Oegopsida

Suborder: -

Family:

Common: Gilchrist’s enope squid Alternate: -

VENTRAL VIEW

Photophores on ventral Emarginated in surface of head arranged in 5-7 clear longitudinal lines

Ventral arms (IV) without suckers

Five photophores Ventral arms ventrally on eye cut away to show purple Mantle, arms and Tips of Arms IV buccal area head covered by with three large small photophores photophores 10 mm

Distinguishing features Hectocotylus • Eye not covered by a transparent membrane, Right or left Arm IV. lens in open contact with seawater. • Funnel-locking cartilage a simple straight Size groove and ridge. 40 mm mantle length. • Characteristic purplish buccal area contrasting with the whitish bases of the arms. Distribution • Mantle, head and arms covered with small Mainly northern parts of West Coast, 200 to 1400 m. photophores visible as small spots. • Photophores on ventral surface of head Similar species arranged in ive to seven clearly deined hoylei: Photophores on ventral surface of longitudinal lines, no photophores between head difuse, not arranged in clear longitudinal lines. these lines. • Arms IV without suckers, two to four (usually siedleckyi: Has one very large and four small three) large photophores covered by black photophores on eyes; Arms IV with suckers distally chromatophores on tips of arms. and without photophores; club with one row of • Eyeball with ive photophores ventrally, anterior hooks. Other Abralia sp. have 5-12 photophores and posterior photophores enlarged. on eyes. • Fin strongly emarginated, lacking posterior lobes and not extending past end of mantle. References Jereb & Roper, 2010; Lipinski, 1983; Nesis, 1987; Club Sanchez, 1988. Two series of hooks and one series of suckers on manus.

368 Phylum: Mollusca Cephalopoda

Quick guide to the Jewel Squids, Genus

Table 4: Comparison of species in the genus Histioteuthis. If unsure of the species use the code “Histio”. H. bonnellii H. macrohista H. meleagroteuthis H. miranda H. reversa Mantle length Much shorter Much shorter Longer than Longer than relative to Shorter than head than head than head head head head length Cartilaginous Small tubercles on Large obvious inconspicuous mantle tubercles on dorsal tubercles on None None None midline of mantle dorsal midline and on Arms I-II of mantle and on Arms I-II Large, elongate photophore Present Present Absent Absent Absent on tips of Arms I-III Inner webbing 50% of arm 50% of arm Less than 15% of Up to 15% of Vestigial between length length arm length arm length Arms I-III Number of photophores 17 16 19-21 16 18 around left eye Number of 6 7 7 7 7 buccal lappets

Examples of the buccal crown in Histioteuthis to illustrate the number of buccal lappets; either six (H. bonnellii, left) or seven lappets (H. macrohista and H. miranda, centre and right panels respectively)

369 Phylum: Mollusca Cephalopoda

Histioteuthis bonnellii (HisBon)

Phylum: Mollusca

Class: Cephalopoda

Order: Oegopsida

Suborder: -

Family:

Common: Ornate/Bonnelli’s jewel squid Alternate: -

LATERAL VIEW

Body covered Mantle shorter than Web extends for 50% of with small head arm length photophores

A large elongated photophore on tips of Arms I-III

Right eye much smaller Single row of large than left photophores on Arms III

Large photophores on 50 mm ventral head and Arms IV 17 photophores around right eye

Distinguishing features Distribution • Eye not covered by a transparent membrane, Mainly on West Coast, from 500 to 1500 m. lens in open contact with seawater. • Body covered with minute photophores. Similar species • Left eye much larger than right. See Table 4 (page 369). • Mantle short, subequal to head length; no The only Histioteuthis sp. in area with six lappets. cartilaginous tubercles on mantle or arms. • Arms joined by an inner web to 50% or more of H. macrohista: Mantle short; buccal lappets seven; arm length. right eye photophores 16; inner web >50% of arms; • Tip of each of Arms I–III bearing a single large ventral surface of head and Arms III and IV without elongate photophore. large compound photophores. • Buccal membrane with six lappets (see image H. meleagroteuthis: Mantle short; buccal lappets on page 369); 17 (rarely 16 or 18) photophores seven; right eye photophores 19-21; inner web around right eye. <15% of arm; single row of large cartilaginous • Large compound photophores on ventral tubercles on dorsal midline of mantle and of surface of head and on Arms III and IV. Arms I-III. Club H. miranda and H. reversa: Mantle long; buccal Small, with four to eight rows of suckers of varying lappets seven. sizes. References Hectocotylus Jereb & Roper, 2010; Nesis, 1987; Sanchez, 1988. Both dorsal arms.

Size Up to 330 mm mantle length (largest Histioteuthis). 370 Phylum: Mollusca Cephalopoda

Histioteuthis macrohista (HisMac)

Phylum: Mollusca

Class: Cephalopoda

Order: Oegopsida

Suborder: -

Family: Histioteuthidae

Common: Plain jewel squid Alternate: -

LATERAL VIEW

Body covered Mantle length less Large photophores on with small than head length tips of Arms I, II & III photophores

Buccal membrane with Large photophores on seven lappets tips of Arms I, II & III

10 mm

Right eye much No large obvious compound 16 photophores around smaller than left photophores on ventral head and Arms III & IV right eye

Distinguishing features Distribution • Eye not covered by a transparent membrane, Both coasts, but more common on West Coast; lens in open contact with seawater. 100 to 1 000 m. • Body covered with minute photophores. • Left eye much larger than right. Similar species • Mantle short, less than head length; no See Table 4 (page 369). cartilaginous tubercles on mantle or arms. H. bonnellii: Mantle short; buccal lappets six; right • Arms joined by an inner web to 50% or more of eye photophores 17; inner web >50% of arms; large arm length. compound photophores on ventral surface of head • Tip of each of Arms I–III bearing a single large and Arms III and IV. elongate photophore. • Buccal membrane with seven lappets (see H. meleagroteuthis: Mantle short; buccal lappets image on page 369); 16 photophores around seven; right eye photophores 19-21; inner web right eye. <15% of arm; single row of large cartilaginous • Ventral surface of head and Arms III and IV plain, tubercles on dorsal midline of mantle and of without large compound photophores. Arms I-III. H. miranda and H. reversa: Mantle long; seven Club buccal lappets. Small, four to eight rows of suckers of varying sizes. References Hectocotylus Jereb & Roper, 2010; Nesis, 1987; Sanchez, 1988. Both dorsal arms.

Size Up to 70 mm mantle length.

371 Phylum: Mollusca Cephalopoda

Histioteuthis meleagroteuthis (HisMel)

Phylum: Mollusca

Class: Cephalopoda

Order: Oegopsida

Suborder: -

Family: Histioteuthidae

Common: Crested jewel squid Alternate: -

LATERAL VIEW Large cartilaginous tubercles Buccal membrane with seven lappets

19-21 photophores 30 transverse rows of around right eye photophores on mantle Eight to nine longitudinal Tentacular rows of photophores at base club of Arms IV

50 mm

Distinguishing features Hectocotylus • Eye not covered by a transparent membrane, Both dorsal arms. lens in open contact with seawater. • Body covered with minute photophores. Size • Left eye much larger than right. Up to 114 mm mantle length. • Mantle short, subequal to head length; inner web between arms less than 15% of arm length. Distribution • No large elongate photophores at the tips of West Coast. Of the shelf in water column to over arms. 1 000 m. • Buccal membrane with seven lappets; 19-21 photophores around right eye. Similar species • Eight to nine longitudinal rows of photophores See Table 4 (page 369). in basal parts of Arms IV; 30 transverse rows of photophores on ventral mantle. Distinguished from other Histioteuthis by large • Large cartilaginous tubercles on dorsal cartilaginous tubercles on dorsal midline of mantle midline of mantle and basal parts of Arms I–III. and dorsal base of Arms I-III; 19-20 photophores around right eye. Club Small, with four to eight rows of suckers of varying References sizes. Jereb & Roper, 2010; Nesis, 1987; Sanchez, 1988.

372 Phylum: Mollusca Cephalopoda

Histioteuthis miranda (HisMir)

Phylum: Mollusca

Class: Cephalopoda

Order: Oegopsida

Suborder: -

Family: Histioteuthidae

Common: Common jewel squid Alternate: -

DORSAL VIEW

Small cartilaginous tubercles

Right eye surrounded by 16 photophores

10 mm Right eye much Web extends for 15% Buccal area showing smaller than left of arm length the restricted webbing between the arms and seven lappets in the buccal membrane

Distinguishing features Size • Eye not covered by a transparent membrane, Up to 270 mm mantle length. lens in open contact with seawater. • Body covered with minute photophores. Distribution • Left eye much larger than right. Most common Histioteuthis species in the region, on • Mantle length greater than head length, with both coasts in 700 to 900 m. small inconspicuous cartilaginous tubercles on dorsal midline of mantle and basal parts of Similar species Arms I–III. See Table 4 (page 369). • Inner web connects basal 15-25% of Arms I-III; outer web not developed. H. bonnellii, H. macrohista and H. meleagroteuthis: • No large elongate photophores at the tips of Mantle length less than head length. arms. H. reversa: Lacks tubercles on dorsal midline and base • Buccal membrane with seven lappets; 16 of arms; 18 photophores around right eye; inner web photophores around right eye. between arms vestigial.

Club References Manus with closely packed suckers of varying sizes Jereb & Roper, 2010; Nesis, 1987; Sanchez, 1988. in six to seven series.

Hectocotylus Both dorsal arms.

373 Phylum: Mollusca Cephalopoda

Histioteuthis reversa (HisRev)

Phylum: Mollusca

Class: Cephalopoda

Order: Oegopsida

Suborder: -

Family: Histioteuthidae

Common: Reverse jewel squid Alternate: -

DORSAL VIEW Left eye much No cartilaginous tubercles bigger than right on arms

Buccal membrane with seven lappets

Inner webbing vestigial

No cartilaginous tubercles on 18 photophores mantle around right eye Tentacular Club

Figure reproduced from Jereb & Roper, 2010, with permission.

Distinguishing features Size • Eye not covered by a transparent membrane, Up to 200 mm mantle length. lens in open contact with seawater. • Body covered with minute photophores. Distribution • Left eye much larger than right. Occurs of Namibia; possible on northern West Coast; • Mantle elongate, much longer than head, 300–1 000 m. lacking cartilaginous tubercles on dorsal midline. Similar species • Arms robust, of moderate length, without See Table 4 (page 369). cartilaginous tubercles or terminal photophores. • Inner web between Arms I-III low, vestigial. H. bonnellii, H. macrohista and H. meleagroteuthis: • Buccal membrane with seven lappets; 18 Mantle length less than head length. photophores around right eye. H. miranda: 16 eye photophores; inner web on Arms I-III <15% of arm; single row of small cartilaginous Club tubercles on dorsal midline of mantle and of Manus with deep longitudinal cleft on aboral surface; Arms I-III. suckers in six diagonal series, median ventral series enlarged (three to four times marginal). References Jereb & Roper, 2010; Nesis, 1987; Sanchez, 1988. Hectocotylus Both dorsal arms.

374 Phylum: Mollusca Cephalopoda

Joubiniteuthis portieri (JouPor)

Phylum: Mollusca

Class: Cephalopoda

Order: Oegopsida

Suborder: -

Family: Joubiniteuthidae

Common: Joubin’s squid Alternate: -

DORSAL VIEW

Arms I-III more than twice mantle length Fins fused, circular

VENTRAL VIEW Arms IV very short Tail longer than mantle

Figure reproduced from Young & Roper, 1969, with permission.

Distinguishing features Size • Eye not covered by a transparent membrane, 105 mm mantle length. lens in open contact with seawater. • Funnel-locking cartilage oval, without knobs. Distribution • Arms I-III very long, more than 2x ML, with very West Coast, very rare. Meso- to bathypelagic from small suckers in six series. 500 m to over 3 000 m. • Arms IV short (1/3 length of other arms), with suckers in four series. Similar species • Head narrow, eyes small without photophores. None. Combination of long Arms I-III and long tail • Mantle long and narrow. diagnostic. • Fin round, short (30% ML); long thin tail (longer than mantle). References Jereb & Roper, 2010; Nesis, 1987; Young & Roper, Club 1969; Sanchez, 1988. Long and laterally compressed; minute suckers in 5-12 transverse series; no carpus.

Hectocotylus Absent.

375 Phylum: Mollusca Cephalopoda

Lycoteuthis lorigera (Lycote)

Phylum: Mollusca

Class: Cephalopoda

Order: Oegopsida

Suborder: -

Family:

Common: Crowned irely squid Alternate: diadema

VENTRAL VIEW

Row of ive light organs on ventral eye margin Luminous bar Transverse row of ive photophores All arms attenuated

Fins translucent, NOT black (photograph Two round photophores was taken on black background) Row of photophores on ventral margin of eye 10 mm

Distinguishing features Club • Eye not covered by a transparent membrane, Four rows of suckers. lens in open contact with seawater. • Large luminous organs in body cavity visible Hectocotylus through the ventral mantle: Two round Absent. photophores near mantle opening; ive round photophores in a transverse row across mantle Size anterior to edge of in; a luminous cross bar near Males 190 mm ML. Females 110 mm. the posterior end of the mantle. • Ventral side of eyeball with ive luminous organs Distribution arranged in a single row. Both West and South Coasts. Deeper than 300 m. • No hooks present. Suckers in two series on arms and four series on clubs. Similar species • Males: Arms II greatly elongated, with a series None. The three series of visceral photophores of regularly spaced photophores; Arms III (visible through the mantle) diagnostic in the area. elongated, strongly attenuated. • Muscular, conical mantle. References • Fins broad, rhomboidal. Jereb & Roper, 2010; Nesis, 1987; Sanchez, 1988.

376 Phylum: Mollusca Cephalopoda

Mastigopsis hjorti (MasHjo)

Phylum: Mollusca

Class: Cephalopoda

Order: Oegopsida

Suborder: -

Family: Mastigoteuthidae

Common: Hjort’s whiplash squid Alternate: hjorti

LATERAL VIEW 2 photophores on ventral eyeball

Diamond-shaped in 90% of ML

Arms IV thicker and longer than other arms Long, worm-like tentacles

Fin width greater than Clubs long and mantle length slender

From Vecchione & Young (2014), reproduced with permission from RE Young

Distinguishing features Size • Eye not covered by a transparent membrane, 100 mm mantle length. lens in open contact with seawater. • Funnel-locking cartilage oval without knobs. Distribution • Mantle weakly muscular, semi-gelatinous. Both West and South Coasts. Oceanic pelagic or • Two photophores on ventral surface of benthopelagic. eyeball; no other photophores on eye or body surface. Similar species • Arm suckers biserial, no hooks; ventral arms Octopoteuthis sicula: Mantle produced posteriorly thicker than other arms, greatly elongated. into a “tail”; armature of hooks; no photophores on • Fins very large, diamond-shaped, reaching eyeball; tentacles residual or absent. almost to anterior edge of mantle (about 90% of ML); width greater than ML. : Tentacles residual; no photophores on eyeball; arms with hooks; large, swollen terminal Club photophore at tips of Arms II. Tentacles vermiform, extremely long, slender; club elongate, with numerous minute suckers arranged References in more than 15 series. Jereb & Roper, 2010; Nesis, 1987; Sanchez, 1988; Vecchione & Young, 2014. Hectocotylus Absent.

377 Phylum: Mollusca Cephalopoda

Octopoteuthis sicula (Octhis)

Phylum: Mollusca

Class: Cephalopoda

Order: Oegopsida

Suborder: -

Family: Octopoteuthidae

Common: Rüppell’s octopus squid Alternate: -

DORSAL VIEW VENTRAL VIEW

Tentacles absent Robust arms or residual bearing hooks

Fin very large

Long tail 50 mm

Distinguishing features Size • Eye not covered by a transparent membrane, 200 mm mantle length. lens in open contact with seawater. • Arms robust, with biserial hooks enveloped Distribution in soft integumentary sheaths. Minute suckers Both West and South Coasts. Meso- to bathypelagic only at tips. down to about 2 000 m. • Small, black, spindle-shaped terminal photophore at the tips of each arm. Similar species • One pair of photophores embedded in posterior Taningia danae: Fin width much greater than ML; mantle; and three pairs on lateral sides of funnel large, swollen terminal photophore at tips of Arms groove near neck. II; no terminal photophores on other arms. • Fin large, length ca 90% ML, width ca 115% ML. Ancistrocheirus lesueurii: Armature of hooks, Club diagnostic brown dashed lines on in. Tentacles present in paralarval stage (up to 15 mm Mastigopsis hjorti: Tentacles present; armature of ML) only, absent in adults. suckers without hooks; two photophores on eyeballs.

Hectocotylus References Absent. Jereb & Roper, 2010; Nesis, 1987; Sanchez, 1988.

378 Phylum: Mollusca Cephalopoda

Taningia danae (TanDan)

Phylum: Mollusca

Class: Cephalopoda

Order: Oegopsida

Suborder: -

Family: Octopoteuthidae

Common: Taning’s octopus squid Alternate: -

DORSAL VIEW

Tentacles absent or residual

Terminal light organ on Fin very large and Arm II broad (© Paul Clerkin)

Distinguishing features Size • Eye not covered by a transparent membrane, 1 700 mm mantle length, 161 kg. lens in open contact with seawater. • Arms robust, with large hooks in two series to Distribution tips. Minute suckers sometimes at tips. Both West and South Coasts. Meso- to bathypelagic • Tips of Arms II with large, oval, swollen down to about 2 000 m. photophore. No photophores on other arms. • No photophores embedded in mantle or arms. Similar species One photophore on either side of intestine Octopoteuthis sicula: Mantle extends as a tail ventral to the ink sac. posterior to broad in; small, black, spindle-shaped • Fin very large, length ca 100% ML, width ca terminal photophore at the tips of each arm. 130% ML. Ancistrocheirus lesueurii: Armature of hooks, Club diagnostic brown dashed lines on in. Tentacles present in paralarval stage (up to 45 mm Mastigopsis hjorti: Tentacles present; armature ML) only, absent in adults. suckers without hooks; two photophores on eyeballs.

Hectocotylus References Absent. Jereb & Roper, 2010; Nesis, 1987; Sanchez, 1988.

379 Phylum: Mollusca Cephalopoda

Ommastrephes bartramii (OmmBar)

Phylum: Mollusca

Class: Cephalopoda

Order: Oegopsida

Suborder: -

Family:

Common: Neon lying squid Alternate: -

MALE: Adult females have large triangular DORSAL VIEW membrane on Arms III Foveola with 5-8 folds

Silvery opalescent stripe Fin rhomboidal, 2-5 side on ventral midline Eyes without very large and pockets photophores broad

Photograph of funnel groove showing the Carpus difuse; foveola and side pockets. Side pockets Dorsal mantle 2-5 knobs and 2-4 smooth can be diicult to see 100 mm darker than ventral suckers on fresh specimens.

Largest medial 4-7 toothed suckers with an enlarged suckers proximal tooth in each quadrant to irst knob

Distinguishing features Hectocotylus • Eye not covered by a transparent membrane, Right or left ventral arm, smooth without suckers. lens in open contact with seawater. • Funnel groove: Foveola with ive to eight Size longitudinal folds; side pockets two to ive ( ) 400 mm ML; ( ) 900 mm ML. (usually three to four), distinct, but can be diicult to see on fresh, wet specimens. Distribution • No ocular, intestinal or large dorsal Oceanic, ofshore of the 200-m isobath where sea photophores; small scattered subcutaneous surface temperature is 10-25 ˚C. Surface to 1 500 m, photophores embedded in the mantle, head but not close to seabed. and ventral arms (not easily visible). • Arms strong, not attenuated, bearing biserial Similar species suckers; swimming keels well-developed. oualaniensis and Sthenoteuthis • In adult females, the ventral membranes of Arms pteropus very similar, distinguished by large III expand into large, triangular lobes. obvious photophore anteriorly on dorsal mantle. • Colour red, dorsal surfaces typically darker Mantle fused to funnel in S. oualaniensis, not fused than ventral; a long, wide, silvery or golden in S. pteropus or other Ommastrephids. See also opalescent strip on ventral midline from mantle and . opening to the level of the ins. • Fins terminal, large, rhomboidal, slightly References attenuated posteriorly; length 40-50% ML; Jereb & Roper, 2010; Nesis, 1987; Roeleveld, 1988; width 60-85% ML; shorter and wider than Sanchez, 1988. Todarodes.

Club Dactylus with four rows of small suckers. Manus with enlarged suckers, largest suckers with four large pointed teeth (one in each quadrant). Carpal- 380 locking apparatus present. Phylum: Mollusca Cephalopoda

Ornithoteuthis sp. (Ornith)

Phylum: Mollusca

Class: Cephalopoda

Order: Oegopsida

Suborder: -

Family: Ommastrephidae

Common: Atlantic and Shiny squids Alternate: -

Two very similar species that can be identiied with certainty to species only by the structure of the FEMALE: DORSAL VIEW hectocotylus. See next page for identiication of males.

Fin large, sharply Head as wide as or broader lanceolate, than mantle opening Foveola with extended 7-12 folds as a long tail Two visceral photophores

Funnel Eye with one ventral photophore 50 mm Image of funnel groove showing the foveola; Enlarged Well-developed side pockets indistinct medial swimming keel and not visible suckers on manus with 18-20 teeth Carpal-locking apparatus weakly developed, lacking knobs

Distinguishing features Hectocotylus • Eye not covered by a transparent membrane, Right Arm IV. Structure difers between species (see lens in open contact with seawater. next page). • Mantle cylindrical and muscular, narrow, extending posteriorly as a long thin tail; head Size broad, equal to or broader than mantle width. 300 mm mantle length. • Fins long, sharply lanceolate, posterior margins concave accentuating the long tail. Distribution • Funnel groove: Foveola with 7-12 folds; side Both West and South Coasts, surface to 1 000 m. pockets obscure few or none. • No external or subcutaneous photophores; Similar species two visceral photophores: one large, round, The Ommastrephid genera are distinguished by the yellowish near the anus, other small, oval, structure of the funnel groove (see Ommastrephes, white at posterior end of intestine, pinkish Todarodes and Todaropsis). See next page for bioluminescent strip extends from the small diferences between the two species in this genus. photophore to posterior tip of mantle cavity. • A single round photophore patch on ventral References surface of each eye. Jereb & Roper, 2010; Nesis, 1987; Roeleveld, 1988; • Arms strong with well-developed swimming Sanchez, 1988. keels; suckers biserial with toothed rings.

Club Suckers in four series. Medial manus suckers very large, with 18-20 equal-sized teeth: carpal-locking apparatus weakly developed, lacking knobs.

381 Phylum: Mollusca Cephalopoda

Ornithoteuthis Males

Phylum: Mollusca

Class: Cephalopoda

Order: Oegopsida

Suborder: -

Family: Ommastrephidae

Common: Atlantic and Shiny bird squids Alternate: -

MALE: DORSAL VIEW

Fin and mantle Males usually skinned extend as long tail in the trawl Hectocotylised right Arm IV

Head as wide as or broader than mantle opening

50 mm

Ornithoteuthis antillarum Ornithoteuthis volatilis Atlantic bird squid Shiny bird squid (OrnAnt) (OrnVol)

Hectocotylus Hectocotylus Ventro-lateral view of right Arm IV Oral and ventro-lateral views of right (igure reproduced from Jereb & Roper, Arm IV (igure reproduced from Jereb & 2010, with permission) showing: Roper, 2010, with permission) showing:

Distal half with sucker stalks modiied Distal half with sucker stalks modiied into papillae; honeycomb sculpturing into papillae; honeycomb sculpturing along midventral surface consisting along midventral surface consisting of four or ive longitudinal columns of two or three longitudinal columns of depressions and swollen ridges. of depressions and swollen ridges. There are 20 to 25 depressions (pits There are 10 to 15 depressions (pits or pores) in each column. or pores) in each column.

Distribution Distribution Known global distribution North Atlantic south to Known global distribution Indo-West Paciic to east at least 28° S of Namibia. coast of Africa. Reported from the Benguela of South Africa and Namibia. Possible of the northern West Coast. Possible of both West and South Coasts.

382 Phylum: Mollusca Cephalopoda

Todarodes angolensis (Toddes)

Phylum: Mollusca

Class: Cephalopoda

Order: Oegopsida

Suborder: -

Family: Ommastrephidae

Common: Angola lying squid Alternate: -

FEMALE: DORSAL VIEW

No external or internal Manus with 14-18 quadriserial Large in with photophores Eyes without rows of suckers short tail photophores Medial manus Carpal area suckers with short, only four 13 to pairs of suckers 16 long, pointed teeth Funnel groove with foveola Club very long, almost Foveola Sketch of funnel groove entire length of tentacle (Figure reproduced from Jereb & Roper, 2010, with permission.) 50 mm

Distinguishing features Size • Eye not covered by a transparent membrane, 430 mm mantle length. lens in open contact with seawater. • Mantle cylindrical and muscular, narrow and Distribution tight in , wider and looser in . Both South and West Coasts. Ofshore of the 300 m • Trawl-caught males usually skinned. isobath. • No light organs on eyes, viscera or mantle. • Arms strong, with well-developed swimming Similar species keels. Suckers biserial, with toothed rings. Ommastrephid genera distinguished by the structure • Funnel groove with foveola (containing of the funnel groove: either smooth (Todaropsis); longitudinal folds) only, side pockets absent. with foveola only (Todarodes) or; with foveola and • Fin large; convex anterior margin; posterior indistinct (Ornithoteuthis) or distinct (Ommastrephes margin attenuated to form short tail. and Sthenoteuthis) side pockets. Club Todarodes ilippovae: Club much shorter; with 12- 14 quadriserial sucker rows; sucker rings with 7-13 Very long; manus with 14-18 quadriserial sucker teeth; carpus very short, only two pairs of carpal rows, medial manus suckers enlarged, with 13-16 suckers; longer in. long pointed teeth. Four pairs of carpal suckers.

Hectocotylus References Jereb & Roper, 2010; Nesis, 1987; Roeleveld, 1988; Right Arm IV long, with suckerless thick pedicels Sanchez, 1988. forming a feather-like fringe for distal 40% of arm.

383 Phylum: Mollusca Cephalopoda

Todarodes ilippovae (TodFil)

Phylum: Mollusca

Class: Cephalopoda

Order: Oegopsida

Suborder: -

Family: Ommastrephidae

Common: Antarctic lying squid Alternate: -

FEMALE: DORSAL VIEW

No external Eyes without photophores Large in with or visceral moderate tail photophores

50 mm

Manus wide, with 12-14 quadriserial rows of suckers

Medial manus suckers with Carpal area short, Arm suckers biserial, with 7-13 long, pointed teeth only two sucker pairs toothed rings

Distinguishing features Size • Eye not covered by a transparent membrane, Max female 540 mm, male 400 mm mantle length. lens in open contact with seawater. • Mantle cylindrical and muscular, narrow. Distribution • No light organs on eyes, viscera or mantle. Circumpolar south of 35° S. Rare on South Coast. • Arms strong, with well-developed swimming Oceanic 300-1 200 m. keels; suckers biserial, with toothed rings. • Funnel groove with foveola only, side pockets Similar species absent. Other Ommastrephids; genera distinguished • Fin large; convex anterior margin; posterior by the structure of the funnel groove (see under margin attenuated to form short tail. T. angolensis). Club Todarodes angolensis: Club much longer, with 14-18 quadriserial sucker rows; sucker rings with 13-16 Short, well-developed; manus wide with 12-14 teeth; four pairs of carpal suckers; shorter in. quadriserial sucker rows; medial manus suckers enlarged, with 7-13 long pointed teeth; carpus very short, only two pairs of carpal suckers. References Jereb & Roper, 2010; Nesis, 1987; Roeleveld, 1988; Hectocotylus Sanchez, 1988. Right Arm IV long, with suckerless thick pedicels forming a feather-like fringe for distal 21-36% of arm.

384 Phylum: Mollusca Cephalopoda

Todaropsis eblanae (Todrop)

Phylum: Mollusca

Class: Cephalopoda

Order: Oegopsida

Suborder: -

Family: Ommastrephidae

Common: Lesser lying squid Alternate: -

FEMALE: MALE: VENTRAL VIEW DORSAL VIEW

Base of hectocotylised ventral arms of Ventral arms mature male slender with with modiied normal suckers suckers

Funnel groove Large eyes without smooth, without photophores foveola or side pockets Four nuchal folds

Manus with six quadriserial rows Large in with convex anterior edge Large medial suckers with 10 mm 30 teeth

Distinguishing features Hectocotylus • Eye not covered by a transparent membrane, Bases of both ventral arms with beak-like lappets, lens in open contact with seawater. edges brown in mature . • Mantle robust, stout, but thinner walled and labbier than Todarodes, especially in . Size • Head broad, with four nuchal folds on neck; 290 mm mantle length in females; 220 mm for males. funnel groove without foveola or side pockets. • Arms strong, with well-developed swimming Distribution keels. Suckers biserial with toothed rings. Both South and West Coasts, 20-850 m. • Largest arm suckers with one large median tooth and three or four smaller teeth. Similar species • No light organs on eyes, viscera or mantle. Distinguished from other Ommastrephids in the • Fin large, broad, width about twice length, area by smooth funnel groove lacking both foveola anterior edge convex. and side pockets; absence of body, eye and visceral photophores; presence of nuchal folds and having Club both ventral arms hectocotylised. Dactylus with four rows of small suckers. Manus with six transverse rows of four suckers, medial suckers References 4x larger than lateral suckers. Largest suckers with Jereb & Roper, 2010; Nesis, 1987; Sanchez, 1988. about 30 teeth.

385 Phylum: Mollusca Cephalopoda

Notonykia africanae (NotAfr)

Phylum: Mollusca

Class: Cephalopoda

Order: Oegopsida

Suborder: -

Family: Onychoteuthidae

Common: Benguela clubhook squid Alternate: -

DORSAL VIEW VENTRAL (See inset): Tentacle stalk triangular V-shaped funnel groove in cross-section Fin large, and mantle margin strongly rhomboidal, emarginated width greater than length Mantle smooth

Arms robust, subequal; suckers biserial 4-5 nuchal folds per side

50 mm Dactylus: 20-38 Ventral view of minute suckers head showing the distinctive V-shaped funnel groove and strongly emarginated Manus: 14-20 strong hooks, Carpal pad ventral mantle no suckers well-deined

Distinguishing features Size • Eye not covered by a transparent membrane, 180 mm mantle length. lens in open contact with seawater. • Mantle densely muscular, broad. Skin smooth, Distribution not rugose. Photophores absent. Common on West Coast. Bathypelagic to 1 200 m. • Nuchal folds four to ive on each side of neck. • V-shaped funnel groove without leshy ridge. Similar species • Arms robust, subequal (33-55% ML) with biserial Todarodes angolensis: Supericially similar, but difers suckers. in the absence of hooks on the clubs, the lack of • Colour maroon to brick red, darker dorsally. V-shaped funnel groove, and ventral mantle margin • Fin large, rhomboidal 58-66% ML, anterior not emarginated. margins slightly convex, posterior almost straight. Onykia robsoni: Skin very rough, “warty”, no photophores; no nuchal folds; long slender tail. Club banksii: Skin smooth; two visceral Dactylus 20-38 minute suckers; manus narrow, two photophores on ventral midline; large light organ medial series of 14-20 (usually 17-18) strong hooks, on eyes; 9-10 pairs of prominent nuchal folds; 20-22 no marginal suckers; carpus well defined, 6-12 large medial hooks on club. smooth suckers plus knobs. References Hectocotylus Jereb & Roper, 2010; Nesis et al., 1998. Absent.

386 Phylum: Mollusca Cephalopoda

Onychoteuthis banksii (OnyBan)

Phylum: Mollusca

Class: Cephalopoda

Order: Oegopsida

Suborder: -

Family: Onychoteuthidae

Common: Common clubhook squid Alternate: -

VENTRAL VIEW DORSAL VIEW

Large patch-like photophore on 9-10 pairs of eye prominent nuchal folds

Two visceral Skin smooth photophores

Tentacular club

Distinguishing features Size • Eye not covered by a transparent membrane, 300 mm mantle length. lens in open contact with seawater. • Mantle very robust, densely muscular. Distribution • Skin smooth, without warts or wrinkles. Possible on both South and West Coasts. Epipelagic, • Nine to ten pairs of prominent, elongate, lap- usually in surface 150 m, but has been recorded to like nuchal folds dorso-laterally on neck. 4 000 m. • A large bi-lobed, patch-like light organ on ventral surface of each eye. Similar species • Two large bulbous visceral photophores on Notonykia africanae: Skin smooth; no photophores; ventral midline, posterior 2x size of anterior. four to ive pairs of nuchal folds; 14-20 medial hooks • Fins moderate, rhomboidal, sharply pointed on club. posteriorly. Onykia species: Skin very rough, “warty”, no Club photophores; no nuchal folds. Dactylus with 13-15 small suckers in four series. Manus slightly expanded with 20-22 large strong References hooks in two medial series; no marginal suckers. Jereb & Roper, 2010; Nesis, 1987; Sanchez, 1988.

Hectocotylus Absent.

387 Phylum: Mollusca Cephalopoda

Onykia robsoni (MorRob)

Phylum: Mollusca

Class: Cephalopoda

Order: Oegopsida

Suborder: -

Family: Onychoteuthidae

Common: Warty squid Alternate: Moroteuthis robsoni

DORSO-LATERAL VIEW

Fin large, Skin very rough attenuated Arms attenuated Long lanceolate tail [similar O. ingens has no tail]

No eye photophores

Very rough Dactylus “warty” skin with minute suckers

Manus with Carpus clearly deined 26-32 hooks 100 mm Detail of mantle skin

Distinguishing features Distribution • Eye not covered by a transparent membrane, Both South and West Coasts in deep waters, 500 to lens in open contact with seawater. 2 500 m. • Mantle robust, long and slender. • Skin rugose, covered with lat, irregular warts. Similar species • Photophores absent on mantle, eyes and Notonykia africanae: Skin smooth; no photophores; viscera; no nuchal folds. four to ive pairs of nuchal folds; 14-20 medial hooks • Arms attenuated with two series of suckers; on club. Arms IV longest. : Skin smooth; two visceral • Fins heart-shaped, very long, attenuated, drawn photophores on ventral midline; large light organ into long lanceolate tail. on eyes; 9-10 pairs of prominent nuchal folds; 20-22 Club large medial hooks on club. Manus long, slender, not expanded, 26-32 hooks : Very similar, but difers in lacking an in two medial series. No marginal suckers. Minute elongated tail, arms not attenuated, and Arms II and suckers on dactylus. Carpus clearly deined. III longer than Arms IV.

Hectocotylus References Absent. Jereb & Roper, 2010; Nesis, 1987; Sanchez, 1988.

Size 900 mm mantle length.

388 Phylum: Mollusca Cephalopoda

Pyroteuthis margaritifera (Pyrote)

Phylum: Mollusca

Class: Cephalopoda

Order: Oegopsida

Suborder: -

Family:

Common: Jewel enope squid Alternate: -

DORSAL VIEW

Clubs with series of three to ive hooks and three series of suckers Fins rounded, subterminal Mantle cone-shaped without photophores Fins not fused posteriorly

End of Arms I-IV long, mantle free two series of of in hooks to tips. Arms without photophores.

Ten photophores in Twelve mantle cavity photophores 10 mm on ventral surface of eye Tentacular stalk with six to seven Figure reproduced from Jereb & Roper, 2010, with permission. photophores

Distinguishing features Hectocotylus • Eye not covered by a transparent membrane, Right ventral arm, without tooth plate. Longitudinal lens in open contact with seawater. membrane along 33% of arm. • Mantle without embedded photophores, cone- shaped. Head broader than mantle. Size • Arms I-IV long and strong, armed with hooks in 50 mm mantle length. two series almost to tips. • Arms without photophores. Distribution • Ventral surface of eye with 12 photophores, nine South and West Coasts. Mesopelagic 400-800 m large and three small. during the day, migrating to upper 200 m at night. • Ten photophores in mantle cavity, three in transverse row at level of the gills. Similar species • Six to seven separated photophores embedded None. in tentacular stalk. • Fins semi-circular, subterminal. References Jereb & Roper, 2010; Nesis, 1987; Sanchez, 1988. Club Manus with a central series of three to ive hooks and two series of suckers.

389 Phylum: Mollusca Cephalopoda

Thysanoteuthis rhombus (ThyRho)

Phylum: Mollusca

Class: Cephalopoda

Order: Oegopsida

Suborder: -

Family: Thysanoteuthidae

Common: Rhombic squid Alternate: -

DORSAL VIEW Photophore on ink sac in young Clubs long, four (less than 350 mm ML) series of suckers Protective membranes on Large rhomboidal long cirri in, widest 1/3 from front

Arms without hooks, two series of suckers

Tentacular club

Funnel-locking 30 mm cartilage shape unique

Figure reproduced from Jereb & Roper, 2010, with permission.

Distinguishing features Hectocotylus • Eye not covered by a transparent membrane, Left ventral arm (IV). Distal third modiied. Small lens in open contact with seawater. untoothed suckers. • Funnel-locking cartilage diagnostic, vertical groove that is joined by a transverse groove Size about halfway along its length in a -shape. 1 300 mm mantle length. • Mantle very muscular and powerful, bluntly rounded. Distribution • Arms short, strong, biserial suckers, no hooks. Of the continental shelf (ofshore of the 400 m • Well-developed protective membranes on isobath) on both South and West Coasts. Pelagic, long cirri-like structures on all arms, but most usually found at or near the surface. obvious on Arms III. • Arms I-III with distinct aboral keels. Similar species • Young squid (60-350 mm ML) with a well- None. developed photophore on ink sac. Reduced, non-functional in adults. References • Rhomboidal, muscular in 100% of mantle Jereb & Roper, 2010; Nesis, 1987; Sanchez, 1988. length, widest 1/3 from front.

Club Tentacles relatively short, strong. Clubs long, widened with four series of suckers. Carpal-locking apparatus a series of alternating knobs and suckers on stalk proximal to clubs.

390 Phylum: Mollusca Cephalopoda

REFERENCES

Adam W and Rees WJ. 1966. A review of the cephalopod family Sepiidae. Scientiic Reports of the John Murray Expedition 1933-1934. British Museum Natural History 11(1): 1-165. Augustyn CJ, Lipinski MR and Roeleveld MAC. 1995. Distribution and abundance of Sepioidea of South Africa. South African Journal of Marine Science 16: 69-83. Jereb P and Roper CFE (Eds). 2005. Cephalopods of the World. An annotated and illustrated catalogue of cephalopod species known to date. Volume 1. Chambered nautiluses and sepioids (Nautilidae, Sepiidae, Sepiolidae, Sepiadariidae, and Spirulidae). FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1. 262pp. 9 Colour plates. Jereb P and Roper CFE (Eds). 2010. Cephalopods of the World. An annotated and illustrated catalogue of cephalopod species known to date. Volume 2. Myopsid and Oegopsid squids. FAO Species Catalogue for Fishery Purposes No. 4, Vol. 2. 605pp. 10 Colour plates. Jereb P, Roper CFE, Norman MD and Finn JK (Eds). 2014. Cephalopods of the World. An annotated and illustrated catalogue of cephalopod species known to date. Volume 3. Octopods and Vampire squids. FAO Species Catalogue for Fishery Purposes No. 4, Vol. 3. 370pp. 11 Colour plates. Lipinski MR. 1983. A description of a new species of enoploteuthid cephalopod, Abralia siedleckyi spec. nov., with some remarks on Abralia redieldi G. Voss, 1955. Veliger, 25(3): 255–265. Lipinski MR. 2001. Preliminary description of two new species of Cephalopods (Cephalopoda: Brachioteuthidae) from South Atlantic and Antarctic waters. Bulletin of the Sea Fisheries Institute, Gdynia 152: 3–14. Nesis KN. 1987. Cephalopods of the World. Multipet, Durban. 351pp. Nesis KN, Roeleveld MAC and Nikitina IV. 1998. A new genus and species of onychoteuthid squid from the . Ruthenica 8:153-168. Roeleveld MA. 1972. A review of the Sepiidae (Cephalopoda) of southern Africa. Annals of the South African Museum 59: 193-313. Roeleveld MA. 1988. Generic interrelationships within the Ommastrephidae (Cephalopoda). In: KM Wilbur (Ed) Paleontology and of cephalopods, Vol. 12, The mollusca. pp. 277–291. London, Academic Press. Roper CFE, Sweeney MJ and Nauen CE. 1984. FAO Species Catalogue. Volume 3. Cephalopods of the World. An annotated and illustrated catalogue of species of interest to isheries. FAO Fishery Synopsis (125) 3: 1-277. Sanchez P. 1988. Systematics and distribution of the cephalopods of Namibia. Monograias de Zoologia Marina 3: 205-266. Sanchez P and Guerra A. 1989. A new species of cirrate octopod Opisthoteuthis vossi from the Southeast Atlantic (Cephalopoda: Octopoda). Bulletin of Marine Science 44: 1159-1165. Young RE. 2009. Vampyroteuthidae Thiele, in Chun, 1915. Vampyroteuthis infernalis Chun, 1903. The Vampire Squid. Ver. 04 July 2012. http://tolweb.org/Vampyroteuthis_infernalis/20084/2012.07.04 in The Tree of Life Web Project, http://tolweb.org/. Young RE and Roper CFE. 1969. A monograph of the Cephalopoda of the North Atlantic: The family Joubiniteuthidae. Smithsonian Contributions to 15: 1-10. Vos NA. 1980. A generic revision of the family Cranchiidae. Bulletin of Marine Science 30(2): 365-412. Voss GL. 1967. Some bathypelagic cephalopods from South African waters. Annals of the South African Museum 50: 61-88 plus 9 plates. Vecchione M and Young RE. 2014. Mastigopsis hjorti (Chun, 1913). Version 06 December 2014. http://tolweb. org/Mastigopsis_horti/19517/2014.12.06 in The Tree of Life Project, http://tolweb.org/. Villanueva R, Collins MA, Sanchez P and Voss NA. 2002. Systematics and distribution in the Atlantic Ocean of the cirrate octopods of the genus Opisthoteuthis (Mollusca, Cephalopoda), with description of two new species. Bulletin of Marine Science 71(2): 933–985.

391 Phylum: Mollusca Cephalopoda

The tube feet of starish leave tiny ‘footprints’ in soft sediments on the South Coast. Photo credit: ACEP Imida Frontiers Project

Brisingid seastars, pumpkin urchin (Dermechinus horridus africanus) and bottlebrush soft corals (Thouarella sp.) at 500 m in the proposed Marine Protected Area on the tip of the Agulhas Bank. Photo credit: ACEP Deep Secrets Project

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