A NEW SPECIES OF SEPIA (CEPHALOPODA: SEPIIDAE) FROM SOUTH AFRICAN WATERS WITH A RE-DESCRIPTION OF SEPIA DUBIA ADAM ET REES, 1966

A new species of cuttlefish Sepia shazae n. sp. is described from South Africa. It is one of the commonest small Sepia species in South African waters occurring from 29°48'S in the north to 25°E in the east, between 200 and 700 m (only the third Sepia species recorded deeper than 600 m). It is recognised by: four papillae clusters dorsally on the head between the eyes; tubercles, warts and prominent clusters dorsally on mantle; skin between these structures smooth and shiny; cuttlebone lightly calcified, thin and fragile with thin inner cone and broad outer cone. S. shazae has been confused with Sepia dubia Adam et Rees, 1966 and is well represented in the holdings of the Iziko Museum, Cape Town (SAMC) as “S. dubia(?)”. S. dubia is re-described here on the basis of the second known individual, and is recognised by: four turret-clusters on dorsal head; two turrets transversely on mid-dorsal mantle; small warts covering dorsal body; cuttlebone heavily calcified, exceptionally broad, especially posterior phragmocone and outer cone. The holotype of S. shazae is deposited in the South African Institute of Aquatic Biodiversity (SAIAB) and paratypes in SAMC and the Natural History Museum (NHMUK), London. The new individual of S. dubia is deposited in SAIAB (the holotype is housed in NHMUK). key Words: small cuttlefish, new species, re-described species, South African waters, Sepia, Hemisepius


INTRODUCTION
Sepia (Sepiidae) and Octopus (Octopodidae) are two exceptionally speciose genera in the class Cephalopoda, a class that is renowned for containing many monotypic genera.Currently Sepia contains 106 nominal species (WorMs 2018).The following species groups of uncertain status (sometimes referred to as sub-genera) have been recognised within Sepia: Sepia s. str., Acanthosepion, Rhombosepion, Anomalosepia, Doratosepion and Hemisepius (khroMov 1998a).
The genus Hemisepius was erected by steenstrup in 1875 for Hemisepius typicus Steenstrup, 1875.Without much explanation adaM & rees (1966) relegated Hemisepius to subgeneric status within Sepia, containing S. (H.) typica and S. (H.) dubia Adam et Rees, 1966. roeLeveLd (1972) discussed the relationship among southern African sepiids in some detail.She retained the subgenus Hemisepius with only two species (S. typica and S. dubia), while noting that S. robsoni (Massy, 1927) and S. faurei Roeleveld, 1972 share a number of characters with Hemisepius and may represent intermediate links "in the transition from Sepia to Hemisepius" (p.257).The discovery of S. pulchra Roeleveld et Liltved, 1985 raised the number of small sepiids in this problematic group to five.roeLeveLd & LiLtved (1985) noted that although these five small sepiids share a number of characters, there are also some worrying morphological differences in important structures such as cuttle-bones and armature of arms.Despite this conclusion, khroMov (1998a) retained Hemisepius as a separate unit containing S. typica, S. dubia, S. pulchra, S. faurei and S. robsoni, with the following characteristics: "Mantle length at maturity <30 mm.Anterior ventral mantle emargination deep, either rectangular or trapezoidal.Dorsal mantle margin slightly produced anteriorly, short, wide.Suckers biserial on all arms.Spine absent.Inner cone markedly reduced, with short limbs." We have participated in numerous demersal surveys conducted in southern African waters by RS Africana  and R/V Dr Fridtjof Nansen (2000)(2001)(2002)(2003)(2004)(2005)(2006)(2007)(2008)(2009)(2010)(2011)(2012)(2013)(2014)(2015)(2016).Extensive new small sepiid material was collected during these surveys and only a fraction of this material has been used in the present paper.It is hoped that this material in its entirety will help to chart research towards resolving the status and relationships between the small sepiids that have traditionally been placed in the so-called Hemisepius complex.These small sepiids appear to form a hitherto unrecognised large group (or groups) of mostly deep water cephalopods, including S. shazae n. sp. a species that is very common and widely distributed in the southern Benguela and is only now recognised as a new to science.

MATERIAL AND METHODS
Specimens of Sepia shazae were collected using bottom trawls during the course of demersal research surveys conducted off the west and south coasts of South Africa by the research vessels RS Africana and R/V Dr Fridtjof Nansen.Details of bottom trawl gear, trawling, sorting and processing of the catch, and references thereof are given in axeLsen & Johnsen (2014), although note that they erroneously record the codend mesh size of the gear deployed by the RS Africana, the correct mesh sizes are 112 mm codend with 35 mm small mesh liner.
Measurements and counts (see: Table 1 for definitions) follow roeLeveLd (1972), roeLeveLd & LiLtved (1985), and Lu & reid (1997) and were taken from preserved specimens.Dorsal (ML) and ventral (MLv) mantle length was measured to the nearest mm below using slide callipers.Fin length (FL) was measured by placing a thread along the base of the fin from the anterior edge and marking the position of the posterior end of the fin, the length of the thread was then measured on a metal ruler.All other measurements were taken using dividers or a graticule in a stereo dissecting microscope at 10× magnification.Sucker diameters were measured at 40× magnification.Weights were taken from preserved specimens.Bodies of most small sepiids in southern African waters have various complex skin patterns that are important for correctly determining species.We therefore developed the following new definitions (or their new combinations) for some of these structures: -Warts are simple, solid, rounded, flat, and low excrescences on the skin, that may be quite large.-Tubercles are simple projections or protuberances, that are sharp or rounded.They may be very long, or short, but never flat.-Papillae are prominent, complex protuberances in following forms: -Turrets (see : roeLeveLd & LiLtved, 1985) where tubercles and/or warts are on top of each other; -Clusters where tubercles and/or warts are next to each other, forming a distinct unit; -Turret-clusters where turrets and clusters are combined in one distinct unit.
Most of the photographs were taken using Canon EOS 7D Mk I and Mk II cameras, or Canon EOS 650 camera coupled with Nikon stereomicroscope using a specially engineered ring.Images of sucker rings and spermatophore were taken using a Nikon SMZ18 stereomicroscope with a P2-SHR Apo1x lens and NIS Elements D 4.60.00(build1171) 64bit software.Images of the radula of S. shazae and S. dubia were taken using a Nikon SMZ1500 stereo dissecting microscope fitted with a Nikon U3 Digital Sight camera system and the Nikon Imaging Systems Basic Research software package.

D i a g n o s i s
Cuttlebone lightly calcified, thin and fragile; anterior part (~30% of length) triangular, pointed; posterior part broad, oval, rounded posteriorly; spine absent; thin chitin flange all round; last septum elevated (occupies ~20% of length); distinct mid-dorsal longitudinal ridge; phragmocone well defined, striae borderline approximately straight or slightly convex; inner cone thin, extends anteriorly into striated zone, Table 3. Minimum (Min), maximum (Max), mean, standard deviation (Std) and sample size (n) for mantle length (ML, mm) and indices (expressed as proportion of ML except for MHL, which is reported as proportion of Hcl) for male and female Sepia shazae sp.nov.examined.The values recorded for the holotype are given for comparison.Individuals for measurements were selected from the whole material available.Main criterion was the biological condition (not all specimens can be measured).See:

D e s c r i p t i o n
Small species; mean±SD ML males 24.4 ±3.84 mm, females 27.6 ±4.45 mm (Table 3).Mantle rather elongated, oval (more round in smaller animals), dorso-anterior margin generally straight wide Ʌ-shape (varies from close to 90° to broad angle, Fig. 2), ventro-anterior margin entire or slightly and broadly emarginated in both sexes (Fig. 3).Ventral margins of mantle with distinct keels (Fig. 3), keels Colour in preservative highly variable, very pale to dark grey or reddish brown (Fig. 2).Skin covered by various structures (papillae, tubercles and warts); smooth, shiny between structures.Skin sculpture (excrescence) species-specific, although quite variable.Four papillae clusters, each with prominent (rarely flattened, unobtrusive) central tubercle dorsally on head between eyes (Figs 4-5); transverse tubercle row between the eyes in occipital region; 3-4 transverse tubercle rows dorsally on head anterior to eyes at arm bases (some rows may be difficult to see).Single small tubercles along arm pairs I-III.Prominent tubercles on flat triangular raised structure anterio-ventral to each eye, ventral tubercle (at triangle tip) largest (Fig. 5).Usually three prominent papillae clusters (sometimes only large, long central tubercles present) dorsally on mantle (Fig. 6), one on dorsal midline near anterior mantle margin, two medially in transverse row midway between dorsal and posterior mantle edges; smaller clusters (usually two pairs) may be present dorsally on posterior mantle.Mantle also covered in mostly irregularly placed tubercles and a few warts.
Tentacular stalk very long (up to ~2.5× ML), club tiny (Fig. 7) with subequal small suckers in eleven diagonal rows of four suckers each.Protective membranes relatively narrow, well separated.Natatory membrane well developed, broad, continuing along tentacular stalk for about 1/3 of club length.
Arms robust, stout, subequal in length (Figs 8-9), relatively long, strong membrane joining pair I proximally for about 40%, membrane becoming gradually shallower between subsequent arms, absent ventrally between arms IV.Protective membranes well developed.Suckers globose, biserial on all arms (sometimes in crowded zigzag pattern appearing tetraserial on arms III and IV, especially in females, considerable variability in this regard among various individuals (see: Remarks).Sucker rings on club (Fig. 11) and arms (Fig. 10) smooth.
Right ventral arm also modified (Fig. 14), sucker arrangement from base to tip: two pairs of biserial normal suckers; two rows of three suckers each; three rows of tetraserial suckers, with marginal suckers of reduced diameter; three pairs of small suckers; 3-4 rows of enlarged biserial suckers; 6-10 rows tiny biserial suckers to tip.Beaks small, fragile, of typical sepiid proportions.Upper beak (Fig. 15): rostrum blunt, relatively short, slightly hooked, length slightly greater than width, rostrum angle curved; hood high above crest poste-riorly; jaw edge straight, jaw angle 90°; lateral wall posterior edge slightly curved; only rostrum and anterior part of hood dark.Lower beak (Figs 16-18): rostrum short, blunt, jaw angle rounded, broad, >90°, cutting edge straight; hood low on crest; crest slightly curved; crest and lower edge of lateral wall at slight angle; posterior edge of lateral wall straight and rounded; only rostrum, anterior part of hood and anterior part of shoulders dark.

R e m a r k s
In size range investigated this small species (largest known specimen is 33 mm ML) is sexually mature at 23-30 mm ML (males), and 20-33 mm ML (females).Juveniles have not been recorded.The holotype (Fig. 1) and most of the paratypes are mature.Observed natural variation and sexual dimorphism that could aid identification in the field has been included in the description.
There is considerable variability in the total number of suckers, especially in females (Table 3).Most of this variability appears to stem from variation in the number of small to minute suckers at the tips of the arms, which could be the result of injury or regeneration.We could not test this in females because the sucker diameter decreases gradually over the length of the arm.Males possess a group of enlarged suckers near the tip of arms II and III (see for example: Fig. 9) that provides a convenient landmark.For a sample of eight males we calculated the absolute  voss 1969: p. 725).In small sepiids, sperm mass is relatively long and other parts relatively short, but not to the extreme shown for example by Australian S. senta Lu et Reid, 1997(Lu & reid 1997: p. 294).
The combination of: small size at maturity (ML seldom larger than 30 mm); anterior mantle margin slightly produced, short and wide; biserial suckers on all arms; lack of posterior spine on the cuttlebone place this species in the so-called Hemisepius-group together with S. dubia, S. faurei, S. pulchra, S. robsoni, and S. typica (part of the Hemisepius diagnosis by khroMov 1998a).

D i a g n o s i s
Cuttlebone well calcified, unusually broad, especially posterior phragmocone and outer cone (relation of length to width ~1.4); anterior triangular; posterior spine absent; dorsal surface covered with calcareous structures, forming reticulate pattern in holotype.Inner cone distinct, broad laterally and posteriorly; together with posterior outer cone forms shallow spoon-shaped depression; limbs fused to outer cone.Outer cone exceptionally broad, surrounds inner cone.Last loculus triangular rather than trapezoid.Striae wavy, almost straight.Head with four turret-clusters dorsally above eyes, two straight transverse rows of warts between eyes, single turret-cluster under each eye.Dorsal mantle: two large prominent turrets close together on either side of dorsal midline near centre of dorsal mantle; at least four smaller turrets anterior and two posterior to large prominent turrets.Many small warts cover dorsal mantle, head and arms.Arms subequal in length, fleshy; strong keels especially prominent in ventral (IV) arms; protective membranes thick and long: suckers biserial, pairs widely spaced, appearing uniserial when laterally squashed (Fig. 31).Arms connected by membrane; slightly more than 1/3 of arm length between pair I; extent gradually decreasing towards arms IV; no web between pair IV.Suckers on arms minute; sucker rings with no teeth (Fig. 33).
Club small; carpus plus manus with 11 rows of five suckers; dactylus with five rows of 2-5 suckers (Fig. 34); suckerless patch in photograph due to damage.Suckers minute, with 4-5 tiny blunt teeth on proximal side of sucker ring only (Fig. 32).
Upper beak (Fig. 35): rostrum blunt, relatively short, not hooked, length equal to width, rostrum angle well defined; hood long, distal tip far from crest, jaw edge straight, jaw angle <90°; lateral wall with posterior edge.Only rostrum and anterior part of hood dark.
Lower beak (Figs 36-38) characteristic: rostrum short, blunt, lacking distinct rostrum angle; hood low on crest, slightly curved, indented; crest slightly curved, not indented, not parallel to lower edge of the lateral wall (proximally further apart than distally); lateral wall with curved and rounded posterior margin.Only rostrum and anterior parts of hood and shoulders dark.
Funnel organ with dorsal component well defined, bearing anterior ridge and papilla, limbs short: ventral component well defined, oval, elongated (Fig. 42).
In discussing the structure of the cuttlebone among species in the Hemisepius group, roeLeveLd & LiLtved (1985, p. 14) state: "Sepia dubia has a Hemisepius-like shell, with the phragmocone having an inverted triangular shape and occupying little more than half the shell length, as in Sepia (Hemisepius) typica."and "Sepia typica, S. dubia and S. faurei have a Hemisepius-like shell with an abbreviated phragmocone, whereas that of S. pulchra is typically sepiid."We found that among these small sepiids, the cuttlebone of S. dubia is the most calcified and solid.In addition it is exceptionally broad, especially across the posterior phragmocone and outer cone and relatively short resulting in a length/width ratio of ~1.4 (which may distinguish it from all other species of Sepia).The shape of phragmocone is also unique and does not resemble that of S. typica (compare our Figs 43-49 with that in roeLeveLd 1972, p. 259 Fig. 17cd).In our view each of these small sepiid species have unique cuttlebone features and there is no "Hemisepius-like cuttlebone".

D i s t r i b u t i o n
Currently known from only two localities, the type locality in False Bay, South Africa and from 119 m depth off the South African south coast at 34°18'00.0"S,24°51'00.0"E(Fig. 27).

DISCUSSION
The small sepiids from southern Africa (S. typica, S. robsoni, S. dubia, S. faurei and S. pulchra) have been grouped as the "Hemisepius species complex" by khroMov (1998a) despite doubts expressed by earlier authors.Being a small sepiid Sepia shazae would also fall into the Hemisepius-group.However the two sepiids described here (S. shazae and S. dubia) are so different from one another that virtually the only trait that they share is their small size, and this trait alone cannot have any systematic significance.If "Hemisepius" were to include the six small sepiids mentioned above, it would unite under one name such contrasting characters as: normally calcified cuttlebone (S. pulchra and S. dubia) vs. not normally calcified or not calcified cuttlebone (S. robsoni, S. faurei, S. typica, S. shazae); tips of dorsal arms finger-like, devoid of suckers (S. faurei and S. robsoni) vs. presence of suckers to the end of all arms (the other four described species); skin on the mantle, head and arms almost smooth with sparse warts (S. robsoni and S. typica) vs. various structures on the skin (all the others).Thus of steenstrup's (1875: p. 468) original characters separating "Hemisepius" from other sepiids only the presence of biserial suckers on all arms is shared among the six quite different small sepiids.Only one character separates his original species (S. typica) from all the others: ventro-marginal deep pores on the mantle.
steenstrup (1875) erected the genus Hemisepius for H. typicus on the basis of three characters: deep pores on the ventral surface of the mantle; poorly developed cuttlebone and biserial suckers on all arms (see : roeLeveLd 1972: roeLeveLd : p. 260). adaM & rees (1966) ) expanded Hemisepius to include their newly described taxon, Sepia (Hemisepius) dubia, then state (p.144) "… if we consider dubia to be a Hemisepius,  roeLeveLd & LiLtved (1985) noted that Sepia pulchra shared characters with the subgenus Hemisepius and Sepia s. str.and that "… this further weakens the distinction between the subgenera Sepia and Hemisepius" (p.14).They discussed the relationships between the five small sepiid species (S. typica, S. robsoni, S. dubia, S. faurei and S. pulchra) and note that although they share some characters, there are also some striking differences.They conclude that "… resolution of the relationships within this group of five species must await collection of further specimens of S. robsoni, S. dubia and S. faurei each of which is known only by the holotype" (p.14-15).
With so many contrasting characters among these small sepiids, Hemisepius as defined by khroMov (1998a) appears to be an artificial grouping of species that is unlikely to be monophyletic.
Sepia shazae is potentially important for future ecological research on sepiids of the region and zoo geographical research on all sepiids globally.It is widespread geographically ranging from the cold Benguela System on the South African west coast to the warm temperate waters of the Agulhas Current System on the south coast.It is also found over a wide depth range from 200 to 700 m (samples collected by bottom trawl), reaching the third greatest depth recorded for any Sepia species in the world (khroMov 1998b, Jereb & roper 2005).This may indicate that its cuttlebone is not physiologically functional as a buoyancy regulator, and is the subject of ongoing research (Fuchs & Lipinski, unpublished).
Relatively simple and basic research (e.g.alpha taxonomy) into the world of small sepiids is ongoing to discover and describe additional small sepiids.With the increase in available material and in the number of taxa described, it may now be opportune to research the morphological, functional and genetic interrelationships among these species of small Sepia occurring off South Africa and to resolve the status of "Hemisepius".In particular, the present description of S. shazae sp.nov.highlights the importance of skin structures (warts, tubercles and papillae) as a morphological character in these analyses.
Aspects of molecular ecology may be especially interesting in the case of common widespread species such as S. shazae.Prior to completing the research for this paper we contributed tissue samples from a number of southern African sepiids, including Sepia shazae (as Sepia sp.A and as Sepia dubia) to a project aimed at developing a phylogenetic tree for the family Sepiidae (see: Lupše et al. 2017, where S. shazae is listed as S. dubia).In their best tree, Lupše et al. ( 2017) link three small sepiid species with Sepia hieronis (Robson, 1924).This link is interesting and should be further explored in future research.The voucher specimens for this project are housed at the Australian Museum in Sydney and SAMC.

Table 1 .
AbbreviationDefinition AL1-AL4 Arm Length: length of the right (rt) or left (lt) arm of each designated (1 to 4) arm pair, measured from the inner base of the most proximal sucker to the tip of the arm.AMH Anterior Mantle to Head: length of anterior projection of the dorsal mantle margin measured along the midline from the anterior-most point of the dorsal mantle to a transverse line joining the posteriormost points of the dorsal mantle margin on either side of the midline AS1-AS4 Arm Sucker diameter: diameter of the largest sucker on the right (rt) or left (lt) arm of each designated (1 to 4) arm pair ASC1-ASC4 Arm Sucker Count: total number of suckers on the right (rt) or left (lt) arm of each designated (1 to 4) arm pair ASl4 Arm Sucker left 4: diameter of the largest sucker on the hectocotylised (left ventral) arm ASl4m Arm Sucker left 4 minimum: diameter of the smallest (modified) sucker on the hectocotylised arm CES Club Edge Suckers: number of suckers along the edge of the club from the basal sucker to the most distal sucker ClRC Club Row Count: number of suckers in a single transverse row across the middle of the tentacular club ClS Club Sucker diameter: diameter of the largest sucker on the tentacular club CS# Club Sucker count: total number of suckers on a tentacular club CTR# Club Transverse Row Number: number of transverse rows of suckers on the tentacular club FFu Free Funnel length: measured from the anterior funnel opening to the dorsal attachment of the funnel to the head FIa Fin Insertion anterior: distance from the anterior mantle margin to the anterior junction of fin and mantle FIp Fin Insertion posterior: distance between the posterior junction of the left and right fins with the mantle Sepia sp.A (LesLie & Lipinski 2018: p. 345) Holotype (Fig.1): SAIAB 205824 -mature male ML 30 mm, TW 6.2 g.R/V Dr Fridtjof Nansen demersal survey 2011401, 06 Feb. 2011, Station 94, 31°06'06.0"S,16°34'06.0"Eto 31°05'12.0"S,16°33'42.0"E,bottom trawl 286-285 m.

FL
Fin Length: measured from anterior to posterior insertion along the curve of the mantle at the base of the fin FuL Funnel Length: measured along the ventral midline from the anterior funnel opening to the posterior margin (the ventral mantle has to be cut to expose the posterior edge of the funnel) FW Fin Width: measured from the lateral edge of the mantle to the free edge of the fin HcL Hectocotylus Length: length of the hectocotylised (left ventral) arm measured from the inner base of the most proximal sucker to the tip of the arm HL Head Length: from the anterior tip of the nuchal cartilage to the anterior edge of the dorsal interbranchial membrane between the dorsal arm pair HW Head Width: the greatest width of the head (generally across the eyes) L Length of the cuttlebone along the mid-line MHL Modified Hectocotylus Length: of the modified (proximal) portion of the hectocotylus measured from the inner base of the most proximal sucker to the inner base of the first normal sucker ML Mantle Length (dorsal): measured along the midline from the anterior edge of the dorsal mantle to the posterior end of the mantle MLv Mantle Length (ventral): measured along the midline from the midpoint of the ventral emargination to the posterior end of the mantle Tcl Tentacular club length: measured from the basal sucker to the tip of the club TL Tentacle Length: measured from the point of emergence from the tentacular sac to the tip of the club TrRC Transverse Row Count: Number of suckers in a single longitudinal row across the tentacular club to 31°02'42.0"S,16°38'36.0"E,bottom trawl 256-254 m; SAIAB 205827 -female ML 29 mm, TW 6.3 g and SAIAB 205822 -female ML 31 mm TW 9.0 g, R/V Dr Fridtjof Nansen demersal survey 2013401, 07 Feb. 2013, Station 116, 30°30'48.0"S,15°25'00.0"Eto 30°29'42.0"S,15°23'36.0"E,bottom trawl 301 m; SAMC A090207 -male ML 30 mm, TW 6.6 g.R/V Dr Fridtjof Nansen demersal survey 2006402, 20 Feb. 2006, Station 1265, 31°46'00.0"S,16°56'00.0"Eto 31°45'30.0"S,16°54'30.0"E,bottom trawl 285-279 m; SAMC A090208 -female ML 28 mm, TW 6.6 g.R/V Dr Fridtjof Nansen demersal survey 2012401, 15 Feb. 2012, Station 102, 31°04'00.0"S,15°58'24.0"Eto 31°03'30.0"S,15°56'36.0"E,bottom trawl 357-354 m.Additional material: NHMUK 20180277 -1 male: 22 mm 2.8 g, 5 females: 19 mm 2.6 g, 21 mm 3.6 g, 26 mm 4.4 g, 27 mm 5.6 g, 29 mm 5.8 g, R/V Dr Fridtjof Nansen demersal survey 2013401, 04 Feb. 2013, Station 99, 31°24'06.0"S,16°41'24.0"Eto 31°25'30.0"S,16°42'00.0"E,bottom trawl 299-302 m (same station as NHMUK 20180276); NHMUK 20180279 -1 male: 26 mm 4.3 g, 2 females: 27 mm 7.1 g, 30 mm 6.2 g, R/V Dr Fridtjof Nansen demersal survey 2008401, 10 Feb. 2008, Station

Fig. 4 .
Fig. 4. Dorsal aspect of the head of the Sepia shazae sp.nov.holotype showing the two pairs of supra-orbital papillae clusters and rows of tubercles across the head and bases of the arms (SAIAB 205824, male ML 30 mm).Scale bar 10 mm

Fig. 20 .
Fig. 20.Spermatophore of Sepia shazae sp.nov.(SAIAB 206659, male ML 28 mm) with an insert showing an enlargement of the cement body, oral connective complex and ejaculatory apparatus

Figs
Figs 23-24.Funnel organ of Sepia shazae sp.nov.(SAIAB 206659, female ML 29 mm): 23 -dorsal component of the funnel organ; 24 -ventral component, cut off from the dorsal component (indicated by the dotted line) typica) or completely (S.robsoni, S. faurei) chitinised; distal third of the dor-sal arms devoid of suckers and finger-like (S. faurei and S. robsoni); the lateral fleshy keel on the ventral mantle pierced by 10-12 pairs of pores (unique to S. typica).Sepia shazae is distinguished from S. dubia by the following characters.Mantle more elongated, length to width ratio >1.5 (Figs 1-3) compared to <1.2 (Figs 28-29).Dorsal mantle in S. shazae bears three prominent papillae clusters, one on the dorsal midline near the anterior mantle margin, the other two close to the midpoint of the dorsal midline and scattered tubercles with a few warts, no turrets (Figs 5-6) whereas S. dubia has two large prominent turrets close to the midpoint of the dorsal midline and a minimum of four smaller turrets anterior and two posterior to those prominent structures and numerous irregular warts (Figs 29-30).There are two supra-orbital papillae clusters, each with a prominent central tubercle and three prominent tubercles on a flat raised structure anterio-ventral to each eye in S. shazae (Figs

Fig. 27 .
Fig. 27.Chart showing the location of all survey trawls where Sepia shazae sp.nov.specimens were recorded (black dots) during bottom trawl research surveys off the coast of South Africa, and the localities for the S. shazae holotype (red triangle) and paratypes (red diamonds) and for the S. dubia holotype (blue triangle) and the only other known specimen (blue circle).The contour lines show the 200 and 500 m isobaths

Fig. 34 .
Fig. 34.Club of Sepia dubia (SAIAB 205830, female ML 22 mm).Bald patch resulted from the damage to the club.Scale bar 1 mm

Table 2 .
Measurements (mm), weight (g) and counts recorded for characters of the holotype and of the smallest, average and largest non-type male and female Sepia shazae sp.nov.examined(selectedfrom the whole material available).See: Table1for a description of the characters Table 1 for a description of the characters ventral part reduced; outer cone very broad, width decreasing anteriorly into upper limbs that end close to anterior tip of cuttlebone.Head with four papillae clusters dorsally between eyes with prominent tubercle in centre of each cluster; transverse tubercle row between eyes; 3-4 transverse tubercle rows dorsally at base of arms.Prominent tubercles on slightly raised triangular patch anterio-ventral of each eye, ventral tubercle largest.Dorsal mantle usually with three prominent clusters, but sometimes only large, long central tubercles present; one cluster on dorsal midline near anterior mantle margin, two medially on either side of midpoint of dorsal midline.Mantle covered in irregularly placed tubercles and few warts; skin between clusters, tubercles and warts smooth, shiny.

Table 4 .
Measurements (mm), weight (g) and counts recorded for characters of the two currently known specimens of Sepia dubia.Values for the holotype from adaM & rees (1966).See: Table1for a description of the characters