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, IX

2 ANNALS OF THE

SOUTH AFRICAN MUSEUM

VOLUME 96

ANNALE VAN DIE

SUID-AFRIKAANSE MUSEUM

BAND 96

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

VOLUME 96 BAND

HE TRUSTEES OF THE DIE TRUSTEES VAN DIE

SOUTH AFRICAN MUSEUM SUID-AFRIKAANSE MUSEUM

CAPE TOWN KAAPSTAD

1985-1988

SET, PRINTED AND BOUND IN THE REPUBLIC OF SOUTH AFRICA

BY THE RUSTICA PRESS (PTY) LTD, NDABENI, CAPE

D2109

LIST OF CONTENTS

Page

AGUIRRE URRETA, M. B.

Aptian ammonites from the Argentinian Austral Basin. The subfamily Helican-

eylinae Hyatt. 1894- (Published’'December 1986:).. 2 ....2622 52-556. 5 eee nee 271

AGuIRRE Urreta, M. B. & KLINGER, H. C.

Upper Barremian Heteroceratinae (Cephalopoda, Ammonoidea) from Patagonia

and Zululand, with comments on the systematics of the subfamily. (Published

DWeccmucilO SOs) hn Ont: Peete” ne es cee eA Aes Seem ow ee S15)

AVERY, G.

Late Holocene use of penguin skins: evidence from a coastal shell midden at Steen-

bras Bay, Liideritz Peninsula, South West Africa-Namibia. (Published June

Soe A Perc se Ee Cs NE bea sera ewe ee ae 55)

DINGLE, R. V.

Turonian, Coniacian, and Santonian Ostracoda from south-east Africa. (Published

i Deresrr al over AST) ay ate aaa ae te oes my eect re fea oe en a 1228

ENpDRODY- YOUNGA, S.

Evidence for the low-altitude origin of the Cape mountain biome derived from the

systematic revision of the genus Colophon Gray (Coleoptera, Lucanidae).

busca Sepicmer 19685). aetna e 5 4 oe ne ene oss aaa ROR assem ee 359

GosLINER, T. M. & LILTVED, W. R.

Aspects of the morphology of the endemic South African Cypraeidae with a dis-

cussion of the evolution of the Cypraeacea and Lamellariacea. (Published

UNELIS tale) OOM) ESRC ire rete era aie ae anal BS SiGe wh Ss} aS eae 67

Hu .iey, P. A. & Krerrt, G.

A zoogeographic analysis of the fishes of the family Myctophidae (Osteichthyes,

Myctophiformes) from the 1979-Sargasso Sea Expedition of R. V. Anton

Dour (eupusnec bepriary 1985.) 0! veces Se odes ae pene sb ohare ie 19

KLINGER, H. C. SEE AGUIRRE URreETA, M. B.

KRreEFFT, G. see HULLEY, P. A.

LILTVED, W. R. see GOSLINER, T. M.

LILTVED, W. R. see ROELEVELD, M. A.

ROELEVELD, M. A. & LILTVED, W. R.

A new species of Sepia (Cephalopoda, Sepiidae) from South Africa. (Published

ic UAT EL S95) eae ee eR are One eel ainda See ume Es megs 1

WILLIAMS, G. C.

Morphology, systematics, and variability of the southern African soft coral Alcy-

onium variabile (J. Stuart Thomson, 1921) (Octocorallia, Alcyoniidae).

AES e de Nira BL OSG aire pence hike ees eee ae ns occ eee Sarge oe 241

Volume 96 is complete in 9 parts.

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BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.

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FiscHerR, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs

Zool. exp. gén. 74: 627-634.

Koun, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.

Ann. Mag. nat. Hist. (13) 2: 309-320.

Koun, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean.

Bull. Bingham oceanogr. Coll. 17 (4): 1-51.

THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. Jn: SCHULTZE, L. Zoologische

und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.

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(continued inside back cover)

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 96 Band

February 1985 Februarie

Part i Deel

0°, y

S B:8.B:8-e

SLouig you 8S

A NEW SPECIES OF SEPIA

(CEPHALOPODA, SEPIIDAE)

FROM SOUTH AFRICA

MARTINA A. ROELEVELD

W. R. LILTVED

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

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Obtainable from the South African Museum, P.O. Box 61, Cape Town 8000

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Printed in South Africa by In Suid-Afrika gedruk deur

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Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

A NEW SPECIES OF SEPIA

(CEPHALOPODA, SEPIIDAE) FROM SOUTH AFRICA

MARTINA A. ROELEVELD

W. R. LILTVED

South African Museum, Cape Town

(With 14 figures and 4 tables)

[Ms accepted 31 July 1984]

ABSTRACT

A new species of the genus Sepia is described from the Atlantic Coast of the Cape Penin-

sula, South Africa. Sepia pulchra sp. nov. is the fifth species of Sepia maturing at a small size

(MLd approx. 20 mm) to be recorded from South African waters. The female animal most

closely resembles S. dubia, which has been placed in the problematic subgenus Hemisepius.

However, the shell of S. pulchra indicates that this species is more closely related to members

of the subgenus Sepia s.s. The use of the ventral keels on the mantle, and the position of

spermatophores on the hectocotylus, are illustrated for the first time in sepiids.

CONTENTS

PAGE

Rae CHOI Ue Inc. 0 Says aa ke <a ee ind Big hah Settee. 1

SS CPPOEDELICIE GES Pi MOVE ees Fete Peet ghee hc ashe ws eS ye ei ane et 2

[SJ RUSTE, OM, Sta eh oie MAS hen ce eg a Pee eee a 2

ROO SIS ee ae ees tried CSI? ag ee fia ee 3

WV CSER PEON erates ie See che reer nies sae Oe Aa ee Sil oe Oh 5)

ERC AKG tree ae eRe A oie REMI fu 2 oad Plse Bsg eS 12

Pxbiiibt Swit he rateCUSPECIES «ar LA fac Gieisain sk Sitoe hss Pee nealele siete 14

BiolacicavouseivalOUuSy -- sane. ee. Wed lek ace ee ees Mee awe ea 15

PaConoOWle(SCMICIMS) nee: aeisont cys = hoy) Sd cvae bye seuss eee He ee aS 18

IRCEC RCN CC SE Eres rs Uae olay ic es sheeals We che ce ee su levees 18

INTRODUCTION

On a series of SCUBA dives off the Atlantic Coast of the Cape Peninsula a

number of small sepiids were collected by one of us (W.R.L.). These specimens

were originally thought to be Sepia dubia, a species known from a single female

specimen (Adam & Rees 1966: 119-120), also from South Africa.

On closer examination the newly acquired specimens were found to differ

from the description of S. dubia in the sucker arrangement on the ventral arms

and the structure of the shell. A re-examination of the holotype of S. dubia

Ann. S. Afr. Mus. 96 (1), 1985: 1-18, 14 figs, 4 tables.

2 ANNALS OF THE SOUTH AFRICAN MUSEUM

showed that the female animals of both S. dubia and the new specimens do not

seem to differ significantly (the male of S. dubia being unknown). The shells,

however, are quite different.

The measurements, indices and abbreviations used in this paper are defined

by Roeleveld (1972: 196). Additional measurements and counts are as follows:

EMLv —effective ventral mantle length, i.e. excluding that part of the

mantle lying outside the limits of the ventral keels; measured

along the ventral midline from the anterior edge of the mantle

to the posterior limit of the keels.

HcL —length of the hectocotylized (left ventral) arm.

MHL —length of the (proximal) modified portion of the hectocotylus,

from the most proximal suckers to the first normal (distal)

suckers; the length of the modified portion is calculated as a per-

centage of the total length of the hectocotolized arm (HcL) and

not of the mantle length.

Club suckers —the total number of suckers on the tentacular club.

Furthermore, an estimate of the maturity stage of each specimen is indi-

cated by a roman numeral I-III (see Table 1). Since the sex, size and maturity

TABLE 1

Definition of maturity stages.

Maturity stage

I: immature

II: maturing

III: mature

Male

Spermatophores absent; testis

small, undeveloped.

Testis and accessory reproductive

glands partially developed, but

spermatophores absent.

Spermatophores present in Need-

ham’s sac, penis and/or’ on

hectocotylus.

Female

Ovary and nidamental glands

small, undeveloped.

Ovary and nidamental glands

partially developed; no eggs

visible in ovary.

Nidamental glands large; fully

developed eggs present in

ovary.

stage affect the morphology (especially the degree of modification of the hecto-

cotylus in males), these conditions are relevant in comparing specimens. A

particular specimen can then conveniently be designated, for example, d II 24,

indicating a mature male with a dorsal mantle length of 24 mm.

Sepia pulchra sp. nov.

Figs 1-14; Tables 2-4

Material

Holotype: SAM-S1036, do III 17, Llandudno, Cape Peninsula, 25 m,

collected by W. R. Liltved, 11 October 1982, SCUBA; deposited in the South

African Museum.

A NEW SPECIES OF SEPIA FROM SOUTH AFRICA 3

Fig. 1. Sepia pulchra sp. nov. Dorsal view of

holotype, SAM-S1036, ¢ III 17.

Scale = 5 mm.

Paratypes: SAM-S822, ¢ III 22, Hottentots Huisie, Cape Peninsula, 15 m.

SAM-S982, @ III 21, Bakoven, Cape Peninsula, 21 m. SAM-S1007, ¢ III 16,

21117, Llandudno, Cape Peninsula, 25m (same data as_ holotype).

SAM—S1029, 6 119, 921121, Llandudno, Cape Peninsula, 26m.

SAM-S1030, @ III 22, Llandudno, Cape Peninsula, 40m. SAM-S1031,

3S 1117, 3 Ill 18, Llandudno, Cape Peninsula, 30 m. SAM-S1032, ¢ III 19,

2 III 22, Llandudno, Cape Peninsula, 30 m. SAM-—S1033, 6 III 20, ¢ III 24,

Llandudno, Cape Peninsula, approx. 50 m. SAM-S1034, ¢ II 15, Hottentots

Huisie, Cape Peninsula, 15 m. SAM-—S1038, @ III approx. 19, Llandudno, Cape

Peninsula, 32 m. SAM-—S1039, 2 III 19, Llandudno, Cape Peninsula, 35 m.

Diagnosis

Mantle with numerous complex dorsal papillae and fleshy ventral keels;

suckers biserial on arms I-III and distally on arms IV, quadriserial proximally

on arms IV; sexual dimorphism involving sucker enlargement on arms II and

sometimes III and modification of sucker shape and size on right arm IV in

males; tentacular club with subequal suckers; shell not calcified, delicate,

broadly oval, with phragmocone extending to anterior margin, with reflexed

inner cone and no posterior spine.

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A NEW SPECIES OF SEPIA FROM SOUTH AFRICA 5)

Description

The animals are small, attaining maturity at a dorsal mantle length of

15-22 mm in males and 19-24 mm in females.

The mantle is broadly oval, its anterior margin convex dorsally and deeply

emarginate ventrally around the funnel. The fins are of variable width, usually

fairly wide, beginning relatively far back (the distance between the anterior ends

of the mantle and fin is 23-33% MLd). Posteriorly the fins are rounded and

separate.

The colour is reddish-brown dorsally on the mantle, head and arms, with

the pigmentation of the mantle extending on to the bases of the fins. Mid-

dorsally on the mantle there is a large oval purplish patch. The entire dorsal

surface of the mantle, head and arms is covered with tubercles of variable size

and complexity. The tubercles occur as three main types: simple tubercles,

complex turrets, and even more complex flat oval tubercles. The larger tubercles

are arranged in a distinct pattern that can be traced in all the specimens,

although not all the tubercles of the pattern are readily distinguishable in every

specimen. The degree of distinction of these large tubercles depends on the

preservation and degree of skin contraction or ‘wartiness’ of the specimen; in

addition, a particular large tubercle may have an overall flat oval appearance in

one specimen and be more turret-like in another.

Fig. 2. Sepia pulchra sp. nov. Ventral view of

holotype, SAM-S1036, ¢ III 17.

Scale = 5 mm.

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A NEW SPECIES OF SEPIA FROM SOUTH AFRICA 7

Fig. 3. Sepia pulchra sp. nov. Diagrammatic

representation of relative sizes and distribution of

the larger tubercles on the dorsal mantle surface.

In most specimens (including the holotype) three larger tubercles form cirri

over each eye. On the mantle the pattern is most readily traced from the dorsal

median patch. The patch bears two large, flat, oval tubercles (Fig. 3, a)

posteriorly and two turrets (Fig. 3, b) anteriorly; four slightly smaller turrets

(Fig. 3, c—f) run along the midline of the patch. The first of these median turrets

(Fig. 3, c) is sometimes enlarged, forming a group of three with the two lateral

turrets (Fig. 3, b). The median line of turrets on the patch is continued ante-

riorly by a single turret (Fig. 3, g) between the patch and the anterior mantle

margin. Immediately posterior to the patch a pair of turrets (Fig. 3, h) straddles

the dorsal midline; a second and somewhat larger pair of turrets (Fig. 3, j)

occurs halfway between the posterior end of the patch and the posterior mantle

margin. The latter pair of turrets (Fig. 3, j) is part of a series of eight tubercles

(Fig. 3, j-m) that are arranged around the patch in a line roughly parallel to the

lateral and posterior margins of the mantle. The six anterior tubercles of this

lateral line are usually flat oval tubercles, whereas the posteriormost pair

(Fig. 3, j) is turret-like. A marginal line of tubercles (Fig. 3, n—s) lies outside the

lateral line. The first two tubercles (Fig. 3, n) of the marginal line lie anterior to

the fins and the remaining tubercles are on the fin bases, alternating in position

with the tubercles of the lateral line. The marginal line usually consists of ten flat

oval tubercles. The spaces between the primary tubercles are filled with smaller

tubercles ranging from simple to fairly complex, with several protrusions. All

the principal tubercles described above are present in the holotype, but the

posteriormost four tubercles (Fig. 3, r—s) of the marginal line are not very

clearly distinguishable.

8 ANNALS OF THE SOUTH AFRICAN MUSEUM

The ventral surface of the mantle, head and arms IV is buff in colour and

forms a sole, made up of a pair of fleshy keels on the mantle and the swollen

under-surfaces of the ventral arms (Fig. 2). The keels do not extend to the pos-

terior end of the mantle, nor to the fin bases laterally; the lateral and posterior

areas between the keels and fin bases are a somewhat lighter shade of the colour

of the dorsal mantle surface.

The arms are short and stout, with well-developed fleshy protective mem-

branes. The upper three pairs of arms are joined by an interbrachial membrane.

Arms I and II are trapezoid in cross-section, with a flattened aboral surface;

arms III and particularly arms IV bear lateral keels.

The suckers are globose, with small apertures. The sucker rings have a

smooth inner edge, without teeth.

In the females, the suckers are biserial on arms I, II and III, though when

the arms are compressed the suckers may give the appearance of being triserially

or even quadriserially arranged on the proximal half of the arm, this compres-

sion most commonly occurring on the lateral arms (II and III). The suckers are

largest near the base of the arm and gradually decrease in size distally. On the

ventral arms of the females the suckers are quadriserially arranged on the proxi-

mal half to two-thirds of the arm but become biserial on the distal part.

In males the left ventral arm is hectocotylized. The proximal two-thirds of

the arm is modified: the oral arm surface is wider than usual and bears numer-

ous transverse folds. The suckers of the modified portion are reduced in size and

form two longitudinal series of suckers on either side of the ridged oral region.

The dorsal suckers clearly pertain to two longitudinal series as the suckers alter-

nate in a zigzag manner. The two ventral series of suckers have merged to form

an almost straight longitudinal row. Distal to the modified portion, the suckers

are biserial.

In the holotype, the hectocotylus bears two pairs of widely separated

suckers proximally. The dorsal and ventral margins of the modified region each

bear eleven alternating lateral and medial suckers. Distally the arm bears seven

to eight pairs of biserial suckers; the first few pairs of biserial suckers are larger

than those in the modified region, but the sucker size decreases towards the arm

tip. In the other males the number of suckers bordering the modified portion of

the hectocotylus varies from ten to fourteen on either side of the ridged region.

The right ventral arm is also modified in males. Proximally there is a group

(usually eight) of large globose suckers, irregularly arranged. Following these

there are two to four medium suckers and two to four minute suckers; these

occur either as a transverse row of four medium-sized suckers followed by two

minute suckers, one on each extreme edge of the oral surface, as in the holo-

type, or as one medium-sized sucker on each extreme edge of the oral arm

surface followed by minute suckers in a transverse row of four. This proximal

group of suckers, which occupies about 40 per cent of the arm length, is fol-

lowed by a pair of highly modified suckers (usually the 15th and 16th from the

arm base). These suckers (Fig. 5) are greatly enlarged and asymmetrically

A NEW SPECIES OF SEPIA FROM SOUTH AFRICA

Fig. 4. Sepia pulchra sp. nov. Hectocotylus and right ventral arm of holotype,

SAM-S1036, ¢ III 17. Scale = 2 mm.

ME:

Fig. 5. Sepia pulchra sp. nov. Detail of modified suckers on

right ventral arm of holotype, SAM-—S1036, ¢ III 17.

Scale = 1 mm.

10 ANNALS OF THE SOUTH AFRICAN MUSEUM

elongate, with the ring proximal to the main body of the sucker, their overall

shape being reminiscent of the pitcher plant (Nepenthes). The modified suckers

are followed by a pair of very small suckers. In contrast, the next pair (usually

the 19th to 20th suckers) is much larger, attaining at least the size of the medium

suckers of the proximal group. Thereafter 24 to 28 suckers gradually reduce in

size to the arm tip and are generally biserially arranged, though contraction of

the arm may sometimes give a quadriserial appearance to the suckers on the

middle of the arm. In one specimen (SAM-S1007, 3d III 16) a second pair

of suckers (the 19th to 20th) is modified as well as the 15th to 16th suckers on

the right ventral arm, though not to the extent of the more proximal pair.

A further form of sexual dimorphism is shown in the enlargement of several

suckers towards the tips of the lateral arms in males. The most marked sucker

enlargement occurs on arms II, where there are usually three (but occasionally

four) pairs of enlarged suckers in the 8th to 12th pairs from the arm base. A ten-

dency towards enlargement of suckers is sometimes also shown on arms III.

Where present, this enlargement affects one to three (usually two) pairs of

suckers in the 10th to 14th rows from the arm base. In the holotype there are

seven enlarged suckers on each arm II; these suckers pertain to the 8th to 11th

pairs on left II and the 9th to 12th pairs on right Il. There is no marked enlarge-

ment of suckers on arms III in the holotype.

Fig. 6. Sepia pulchra sp. nov. Dorsal and lateral arms of holotype, SAM-—S1036, ¢ III 17,

showing enlargement of distal suckers on arms II (brackets). Scale = 5 mm.

A NEW SPECIES OF SEPIA FROM SOUTH AFRICA ot

The tentacular club is short and broad, somewhat recurved, and bears sub-

equal suckers in transverse rows of four to six. The dorsal protective membrane

is well developed and separate proximally from the ventral protective mem-

brane, which curves around the base of the club. The natatory membrane is

broad and continues along the tentacular stalk beyond the proximal end of the

club. A terminal pad found on the club of many Sepia species, e.g. Sepia faurei

(Roeleveld 1972, fig. 16b) and S. robsoni (Adam & Rees 1966, pl. 46, fig. 279),

is situated at the anterior end of the dorsal protective membrane and is of

similar size and shape to the club suckers. The dorsal surface of the club bears

several transverse rows and a number of scattered chromatophores.

The beaks, radula and spermatophore are illustrated in Figures 8 and 9.

The lower beak of SAM-—S1033, ¢ III 24, has the following dimensions (after

Clarke 1962 and Wolff 1982): rostral length (RL) 0,86 mm, wing length (WL)

3,24 mm, rostral tip to inner margin of wing (RW) 4,08 mm, crest length (CL)

3,26 mm, hood length 1,50 mm, jaw-angle width (JW) 1,66 mm. The hood and

lateral wall are darkened around the rostrum and jaw angle, but leave a clear

strip between them along the jaw edge. The darkening of the lateral wall

gradually diminishes posteriorly and the wing is only slightly pigmented. The

jaw angle is indistinct and obtuse, and is not obscured by a wing fold. The

lateral wall has no distinct ridge or fold and there is no indentation in its poste-

rior margin. The wing is long in relation to the rostrum (WL/RL 3,8).

Fig. 7. Sepia pulchra sp. nov. Tentacular clubs, SAM-S822, ¢ III 22.

Scale = 1 mm.

12 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 4

Sepia pulchra sp. nov. Indices for shells.

Holotype

SAM-S822 SAM-S982 SAM-S1038 SAM-S1039 SAM-S1036 n Mean Range

3 III 22 ? Il 21 Spike ds 2 Ill 19 3 Ill 17

Lin mm 21 19 19 CoN) 16 16-21

W 52,4 63,2 63,2 Gye 9s 5653 5 ¢. 58,6 52,4=6582

Str z Caled O, SUE) 63,2 Oe SITS 62,5 5. ¢. 61D: 6 S226-e.gtes

The shell is not calcified and is broadly oval, becoming somewhat angular

anteriorly; posteriorly the shell is broad, with a strong ventral curvature and a

dorsal hump over the posterior end of the phragmocone. There is no posterior

spine. The dorsal surface of the shell is finely reticulate and shows no distinct

median ridge; the striae of the phragmocone are clearly visible through the thin

dorsal shield. The phragmocone covers virtually the entire length of the shell

and is very thin; the ventral surface is flat or somewhat concave, with at most an

indistinct median groove; the striated zone is long and triangular, the striae

wavy with an overall convex shape. The inner cone is reflexed and fused to the

outer cone; in the holotype it is narrow, the limbs of the inner cone extending

anteriorly along the lateral edges of the striated zone; in a larger shell

(SAM-S1038, Fig. 9A) the inner cone is rather broader laterally, then tapers

rapidly and twists, curving over the lateral edge of the striated zone. The outer

cone is very broad laterally but narrow posteriorly, separating the posterior end

of the inner cone from the shell edge by a narrow strip of outer cone.

Remarks

The female animals of Sepia pulchra are difficult to distinguish from

S. dubia. Adam & Rees (1966: 120) described the arm suckers of S. dubia as

being biserial, but re-examination of the holotype, 2 II 17, showed that, as in

Fig. 8. Sepia pulchra sp. nov. Beaks, SAM-S1033, ? III 24. A. Upper beak, rostral length

1,0 mm. B. Lower beak, rostral length 0,86 mm.

A NEW SPECIES OF SEPIA FROM SOUTH AFRICA 13

Fig. 9. Sepia pulchra sp. nov. A-B. SAM-S1038, ° III approx. 19. A. Detail of posterior

part of shell, ventral view. B. One row of teeth from radula. C. SAM-—S822, ¢ III 22.

Spermatophore from Needham’s sac, with enlargement of the oral end. Each scale = 0,5 mm.

S. pulchra, the suckers are biserial on arms I-III, with a tendency to irregularity

on the lateral arms due to crowding, and quadriserial in the middle of arms IV.

According to Adam & Rees (1966: 119), in the holotype of S. dubia ‘the

dorsal surface of mantle, head and arms is covered with well spaced, round

papillae, creating a very rugose appearance. .. . On the dorsal surface of the

mantle, approximately in the middle, there are two oval patches of contracted

papillae, one on each side of the median line, and a third one anteriorly near the

mantle-margin.’

On careful scrutiny, the arrangement of the tubercles of the holotype of

S. dubia was found to be remarkably like that of S. pulchra. The oval patch on

either side of the midline in S. dubia corresponds with the two main complex

tubercles (Fig. 3, a) in S. pulchra; the third anterior patch of S$. dubia is much

larger than the corresponding tubercle, c, in S. pulchra and may represent an

amalgamation of tubercles c and d of S. pulchra, as in S. dubia there is only this

one large tubercle between the two smaller tubercles, g and e, found in both

14 ANNALS OF THE SOUTH AFRICAN MUSEUM

species. The area of the median tubercles (a—f) does not appear as a distinct

dorsal patch in S. dubia.

The remaining tubercles (h—-s) on the dorsal mantle surface of S. dubia,

though not very distinct, appear to show the same arrangement as in S. pulchra,

with the exception of tubercle n. In S$. dubia tubercle n lies immediately above,

rather than in front of, the anterior fin margin, since the distance between the

anterior margins of the mantle and the fin is less (12,5 % MLd) in S. dubia than

in S. pulchra (23-33 % ML4d).

At this stage it is not possible to assess the significance of the differences in

the tubercles (c, d, n) and the presence or absence of the median patch in separ-

ating the two species, since S$. dubia is known from a single female specimen.

However, the two species differ markedly in their shells. 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. The shell of S. pulchra, on the other hand, has an overall resemblance to

that of S. tuberculata, though being much more fragile; this further weakens the

distinction between the subgenera Sepia and Hemisepius.

AFFINITIES WITH RELATED SPECIES

Sepia pulchra is the fifth small species of Sepia to be found in South African

waters and further compounds, rather than resolves, the intriguing problem of

their interrelationships. The other species are Sepia robsoni, S. faurei, Sepia

(Hemisepius) typica and S. (Hemisepius) dubia. All five species are small,

maturing at a dorsal mantle length of about 20 mm. They all have a very broad

mantle (MW 60-90% ML4d), the anterior margin of which is convex dorsally and

deeply emarginate ventrally, and fleshy keels ventrally on the mantle; suckers that

are biserial, at least on arms I, II and III; and a tentacular club bearing small sub-

equal suckers and protective membranes that are separate proximally.

Sepia typica, S. dubia and S. faurei have a Hemisepius-like shell with an

abbreviated phragmocone, whereas that of S. pulchra is typically sepiid. The

shell of S. robsoni is virtually unknown. Massy (1927: 160) mentioned only that

‘The calcareous portion of the shell has unfortunately been totally dissolved,

only the membranous part remaining’. Adam & Rees (1966: 121) found the shell

to be in poor condition and compared it to S. hieronis and S. insignis. If the

phragmocone of S. robsoni was not of the normal sepiid type, Adam & Rees

would certainly have mentioned it.

The convenient separation of these five species into three with a hemisepiid

shell and two with a normal shell is not borne out by other characters of the

animals. Sepia robsoni and S. faurei both have dorsal arms with finger-like tips,

devoid of suckers, whereas S. dubia and S. pulchra are virtually indistinguish-

able at present except by the shell characters.

The resolution of the relationships within this group of five species must

await the collection of further specimens of S. robsoni, S. dubia and S. faurei,

A NEW SPECIES OF SEPIA FROM SOUTH AFRICA LS

Fig. 10. Sepia pulchra sp. nov. Fig. 11. Sepia pulchra sp. nov.

Dorsal view of shell of holotype, Ventral view of shell of holotype,

SAM-S1036, ¢ III 17. SAM-S1036, ¢ III 17.

Scale = 5 mm. Scale = 5 mm.

each of which is known only by the holotype. The problem is further compli-

cated by the small size of the animals and the difficulty in extracting the very

delicate shells.

BIOLOGICAL OBSERVATIONS

The collection of live specimens of Sepia pulchra has provided the oppor-

tunity to add a few notes on the biology of the species. The sole, formed by the

fleshy keels on the mantle (Fig. 12) and the swollen under-surfaces of the ventral

arms, is found in several species (see above), but its use has not been figured

before. Since most of the specimens of S. pulchra were observed (by W.R.L.)

adhering to vertical rock faces in the head-down position, the sole must be of

considerable importance in maintaining this position. Camouflage against the

background would then be effected by the extensive tuberculation of the dorsal

surface of the mantle, head and arms; the well-known ability of sepiids to

undergo extreme variation in colour pattern would be a further contributing

factor. The reason for the pigmentation of the part of the ventral mantle surface

between the keels and fin bases in this species became obvious on observing

animals in an aquarium. When an animal adheres to the hard substrate, these

parts of the mantle are clearly visible (Fig. 12) and thus also require to be

camouflaged, whereas in species that settle on a sandy substrate this part of the

mantle is not usually visible.

Two mature males (SAM-S1031) were found to have spermatophores

attached to the hectocotylus, allowing the position of the spermatophores to be

16 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 12. Sepia pulchra sp. nov., adhering to rock by fleshy keels of

mantle; in this instance the ventral arms are not participating in the

~adhesion; SAM-S1039, 2 III 19, photographed in aquarium.

illustrated for the first time (as far as could be established). While copulation has

been observed in S. officinalis (Grimpe 1926; Bott 1938; Tinbergen 1939), it has

been found difficult to see the actual transfer of spermatophores, since this is

obscured by the intermingled arms of the mating animals. Bott (1938: 155) has

given a detailed description of spermatophore transfer from male to female: the

modified (proximal) portion of the hectocotylus, which bears a number of trans-

verse folds on the oral surface bordered by a reduced number of rudimentary

suckers, forms a groove connecting the funnel of the male with the bursa copula-

trix on the buccal membrane of the female. According to Bott’s observations the

hectocotylus thus functions as a passage for the spermatophores.

The structure of the hectocotylus of S. pulchra is basically like that of

S. officinalis. However, observations on S. pulchra suggest that the spermato-

phores not only pass along the hectocotylus but are actually attached to the arm

(Figs 13-14). Furthermore, Figure 14 shows that the spermatophores are

attached to the hectocotylus not only in the ridged area between the rudimen-

tary suckers, but also on the lateral part of the arm outside the sucker rows.

Though the attachment of spermatophores to the hectocotylus is not very firm,

since the spermatophores are fairly easily dislodged in preserved specimens, the

strength of attachment was nevertheless sufficient to maintain the spermato-

phore position during capture, transport and preservation of these two

specimens. The means of attachment of the spermatophores to the hectocotylus

would appear to be mucous, since there was no evidence of any structural

attachment, nor did the suckers appear to play a part.

Examination of the bundle of spermatophores of S$. pulchra illustrated in

Figure 13 showed the spermatophores to have discharged and be interspersed

A NEW SPECIES OF SEPIA FROM SOUTH AFRICA

Fig. 13. Sepia pulchra sp. nov. Position of spermatophore bundle

on modified portion of hectocotylized left ventral arm; SAM-—S1031,

3 III 17. Scale =2 mm.

Fig. 14. Sepia pulchra sp. nov. Position of individual spermato-

phores on hectocotylus, after most of spermatophore bundle

removed; SAM-—S1031, d III 18. Scale = 2 mm.

18 ANNALS OF THE SOUTH AFRICAN MUSEUM

with sperm reservoirs, the bundle being held together by mucus or perhaps

cement from the cement bodies of the spermatophores. The spermatophores in

this bundle were all held together by their oral ends, an apparently unnatural

condition possibly attributable to the capture and handling of the animal.

However, the spermatophores illustrated in Figure 14, which had also ejacu-

lated, were attached to the hectocotylus by their aboral ends, an orientation

more consistent with the observation that spermatophores leave the penis

aboral-end first and are carried to the female oral-end first, being held in posi-

tion by the male until the spermatophores have discharged and the sperm

reservoirs are fixed by the cement to the female, either in the mantle cavity or

on the buccal membrane (Drew 1919: 398, 413).

ACKNOWLEDGEMENTS

We would like to thank Mr F. Naggs of the British Museum (Natural

History) for making the holotype of Sepia dubia available for re-examination.

We would also like to thank the following colleagues at the South African

Museum for their help: Dr V. B. Whitehead for comments on the manuscript;

Mr S. X. Kannemeyer and Ms S. Dove for assistance with the photography.

REFERENCES

ApaM, W. & REES, W. J. 1966. A review of the cephalopod family Sepiidae. Scient. Rep. John

Murray Exped. 11(1): 1-165. ;

Bott, R. 1938. Kopula und Eiablage von Sepia officinalis L. Z. Morph. Okol. Tiere 34:

150-160.

CLARKE, M. R. 1962. The identification of cephalopod ‘beaks’ and the relationship between

beak size and total body weight. Bull. Br. Mus. nat. Hist. (Zool.) 8: 421-480.

Drew, G. A. 1919. Sexual activities of the squid Loligo pealii (Les.). Il. The spermatophore;

its structure, ejaculation and formation. J. Morph. 32: 379-435.

GrimPe, G. 1926. Biologische Beobachtungen an Sepia officinalis. Verh. dt. zool. Ges. 31:

148-153.

Massy, A. L. 1927. The Cephalopoda of the South African Museum. Ann. S. Afr. Mus. 25:

151-167.

ROELEVELD, M. A. 1972. A review of the Sepiidae (Cephalopoda) of southern Africa. Ann. S.

Afr. Mus. 59: 193-313.

TINBERGEN, L. 1939. Zur Fortpflanzungsethologie von Sepia officinalis L. Archs néerl. Zool. 3:

323-364.

Wo rr, G. A. 1982. A beak key for eight eastern tropical Pacific cephalopod species with

relationships between their beak dimensions and size. Fishery Bull. natn. ocean. atmos.

Adm. 80: 357-370.

6. SYSTEMATIC papers must conform to the Jnternational code of zoological nomenclature

(particularly Articles 22 and 51).

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be

followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb.

nov., syn. nov., etc.

An author’s name when cited must follow the name of the taxon without intervening

punctuation and not be abbreviated; if the year is added, a comma must separate author’s

name and year. The author’s name (and date, if cited) must be placed in parentheses if a

species or subspecies is transferred from its original genus. The name of a subsequent user of

a scientific name must be separated from the scientific name by a colon.

Synonymy arrangement should be according to chronology of names, i.e. all published

scientific names by which the species previously has been designated are listed in chronological

order, with all references to that name following in chronological order, e.g.:

Family Nuculanidae

Nuculana (Lembulus) bicuspidata (Gould, 1845)

Figs 14-15A

Nucula (Leda) bicuspidata Gould, 1845: 37.

Leda plicifera A. Adams, 1856: 50.

Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).

Nucula largillierti Philippi, 1861: 87.

Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

Note punctuation in the above example:

comma separates author’s name and year

semicolon separates more than one reference by the same author

full stop separates references by different authors

figures of plates are enclosed in parentheses to distinguish them from text-figures

dash, not comma, separates consecutive numbers

Synonymy arrangement according to chronology of bibliographic references, whereby

the year is placed in front of each entry, and the synonym repeated in full for each entry, is

not acceptable.

In describing new species, one specimen must be designated as the holotype; other speci-

mens mentioned in the original description are to be designated paratypes; additional material

not regarded as paratypes should be listed separately. The complete data (registration number,

depository, description of specimen, locality, collector, date) of the holotype and paratypes

must be recorded, e.g.:

Holotype

SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach

Port Elizabeth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973.

Note standard form of writing South African Museum registration numbers and date.

7. SPECIAL HOUSE RULES

Capital initial letters

(a) The Figures, Maps and Tables of the paper when referred to in the text

e.g. *... the Figure depicting C. namacolus ...’; *. . . in C. namacolus (Fig. 10)...’

(b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded

by initials or full names

e.g. DuToit but A.L.du Toit; Von Huene but F. von Huene

e.g. Therocephalia, but therocephalian

Punctuation should be loose, omitting all not strictly necessary

Reference to the author should be expressed in the third person

Roman numerals should be converted to arabic, except when forming part of the title of a

book or article, such as

‘Revision of the Crustacea. Part VIII. The Amphipoda.’

Specific name must not stand alone, but be preceded by the generic name or its abbreviation

to initial capital letter, provided the same generic name is used consecutively.

Name of new genus or species is not to be included in the title: it should be included in the

abstract, counter to Recommendation 23 of the Code, to meet the requirements of

Biological Abstracts.

MARTINA A. ROELEVELD

W. R. LILTVED

A NEW SPECIES OF SEPIA

(CEPHALOPODA, SEPIIDAE)

FROM SOUTH AFRICA

OF THE SOUTH AFRICAN >

CAPE TOWN

CO ESE ie