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ANNALS OF THE ANNALE VAN DIE

~ SOUTH AFRICAN MUSEUM SUID-AFRIKAANSE MUSEUM

VOLUME 93 BAND 93

ARTHSONI AR

NOV - 9 1984

LIBRARIES 7

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

VOLUME 93 BAND

THE TRUSTEES OF THE DIE TRUSTEES VAN DIE

SOUTH AFRICAN MUSEUM SUID-AFRIKAANSE MUSEUM

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1984

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LIST OF CONTENTS

DE Decker, A. H. B.

Near-surface copepod distribution in the south-western Indian and south-eastern

Niantic Oceana (kublished une lO S4e ewe am gee ce foe misc asa. 6 a.

DINGLE, R. V.

Mid-Cretaceous Ostracoda from southern Africa and the Falkland Plateau.

(RublishedicbnuanyelO Sse ern Crevice Set ee tk Mier tee nes aauNa saa

GOSLINER, T. M. see Marcus, E. D. B.-R.

HuLtey, P. A.

The South African Museum’s Meiring Naude cruises. Part 14. Family Myctophidae

(Osteichthyes, Myctophiformes). (Published February 1984.).................

KENSLEY, B.

The South African Museum’s Meiring Naude cruises. Part 15. Marine Isopoda of the

LO PASTS eT chuisesa (aublishedeune T9844") baer er eer eer a

Marcus, E. D. B.-R. & GOSLINER, T. M.

Review of the family Pleurobranchaeidae (Mollusca, Opisthobranchia). (Published

RCWIUIAy eI SA) res eee pea iree en tee iar Ap rim ames ie san eee Mate a kee ss

Volume 93 is complete in 5 parts.

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ANB OROM TALON SSIS ES ToS oo pas lao id a chee, ots ee eke See a) career 27

Hemingwayella (Parahemingwayella) Dingle, 1984 .............. 0 ccc cece eee 135

IMCL TIGL DOA cg TTC ices 5 te tahoe Cen ae oodles ALOR ane ee ee 166

INGIGIIGNINGINCNSICY PALO SA we wegen es ean Sees ore gle ble cesta ok NS OAR GR Bale ne 279

Pleurobranchaea (Macfarlandaea) Marcus & Gosliner, 1984 ....................02005. 40

POnCOlAGV UCL DIN GIO MIO SA yar ee dere eye aa oe oe ks Dees thor eee hs ba oA 148

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

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

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

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

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

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(continued inside back cover)

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 93 Band

February 1984 Februarie

Part 1 Deel

REVIEW OF THE FAMILY

PLEUROBRANCHAEIDAE

(MOLLUSCA, OPISTHOBRANCHIA)

EVELINE d. B.-R. MARCUS

TERRENCE M. GOSLINER

Cape Town Kaapstad

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Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

REVIEW OF THE FAMILY PLEUROBRANCHAEIDAE

(MOLLUSCA, OPISTHOBRANCHIA)

EVELINE D. B.-R. MARCUS

Department of Zoology, University of Sado Paulo

TERRENCE M. GOSLINER*

South African Museum, Cape Town

(With 25 figures)

[MS accepted 28 June 1983]

ABSTRACT

A review of the Pleurobranchaeidae is provided, including keys to the notaspidean genera

and the described species of Pleurobranchaeidae. Eleven members of the family are considered

as junior synonyms while eight additional species are insufficiently described and their status re-

mains uncertain. The morphology of eleven previously described and six new species is charac-

terized and discussed. New taxa are Pleurobranchaea (Macfarlandaea) subgen. nov., and

Pleurobranchaea confusa, P. augusta, P. bonnieae, P. bubala, P. notmec, and P. vayssierei spp.

nov.

CONTENTS

PAGE

MHOC CEO gery ete ene reese ten me ta BA Eco 1

General characters of the Pleurobranchaeidae.................... 2

igleuropranchaca Ieeles LOIS win scc., se Wok ete is = Bes a ei ees 8

Pleurobranchaea (Pleurobranchaea) Leue, 1813.................. 8

Pleurobranchaea (Macfarlandaea) subgen. nov. .................. 40

EUSCICH OPS MAS OIRO SIO oh lan mses Goch eis ee tae ies mse mics gee 42

leuropranchellatnicle. W925 eiy eis oeyni t+ ees cee ee a elk 43

INGONSIGIN Clie LSS vee os, ce. ye ae ee ha ancl as 46

Gigantonotum Lin Guangyu & Tchang Si, 1965 .................. 48

Zoogeography of the Pleurobranchaeidae ....................... 48

PNCKMOWIECOCINENUS past eras re oe as ays ae moda hae a ns 49

FRETC HE TCC Seppe eae ts ee ee Ti ME ee Sie Cs Sars or 49

INTRODUCTION

This paper was originally intended to be the eighth contribution to the cata-

logue of the western Atlantic warm-water Opisthobranchia (Marcus, Ev. 1972a,

1972b, 1973, 1974, 1978, 1980, 1982a). During the course of investigations it was

* Present address: Dept. of Invertebrate Biology and Paleontology, California Academy of

Sciences, San Francisco.

Ann. S. Afr. Mus. 93 (1), 1984: 1-52, 25 figs.

2 ANNALS OF THE SOUTH AFRICAN MUSEUM

determined that the family Pleurobranchaeidae was in need of substantial re-

vision and the scope of this work was expanded.

The classification of the Pleurobranchaeidae has in many cases been based

upon insufficiently described material, and the species were often simply iden-

tified with previous descriptions from geographically related regions. Vayssiére

(1885: 108) found that the descriptions and diagnoses of the Pleurobranchacea

had been so incomplete that the species were confused, considered as synonyms,

or recorded only as varieties. He tried to define clearly the important characters.

The external morphology is influenced by preservation, contraction, and

loss of pigment. The presence of well-separated rhinophores serves as a familial

distinction.

The jaw plates and radula can be used for generic separation but hardly dif-

fer within a genus. Vayssiére (1901: 72) maintained that jaws and radula are in-

dispensable for classification. He stated that it was a pity that Bergh had not

figured jaws and radula, as the comparison of these organs might have elimi-

nated all doubts (Vayssiére 1901: 51, 71).

Few authors have described the reproductive organs completely, although

these offer good specific characters (Vayssiére 1901: 53). This is probably largely

due to the fact that the reproductive organs are very difficult to dissect without

tearing the various ducts, as indicated by Vayssiére (1901: 38). For this study

there are sufficient specimens of some species, but in the case of several others

complete descriptions cannot be given nor can all specimens be classified. Often

the male organs have not been observed in detail and they are often species-

specific in their structure. For these reasons the present authors have concen-

trated on the determination of penial morphology but describe other characters

whenever possible.

Vayssiére (1898: 234) described the oviduct entering the albumen gland in

the Berthellinae: Berthellina brocki (fig. 180) and Berthella ocellata (fig. 182). In

Berthellina edwardsi (fig. 179) the duct, though touching the albumen gland,

seems to continue (p. 234) to its own outer opening. Vayssiére stated that in the

Pleurobranchidae, Pleurobranchus (figs 177-178), Susania, and Oscanius, and in

the Pleurobranchaeidae there is no such connection; the oviduct goes directly

into the vagina and the gland mass opens into the outer part of the vagina. In

earlier papers the senior author also misinterpreted the close proximity of the

oviduct to the gland mass as an entrance in Pleurobranchus aerolatus (Marcus,

Ev. & Marcus 1967a: 165, fig. 19, 0), Pleurobranchaea gela (Marcus, Ev. &

Marcus 1966: 176, fig. 37, oi), and P. agassizii (Marcus, Ev. & Marcus 1967a:

49, fig. 56D, ov).

GENERAL CHARACTERS OF THE PLEUROBRANCHAEIDAE

The colours are variable in life, but most species are brownish. A shell is

absent. The size differs between species but depends on age and may vary be-

tween populations. The notum varies in size. It is usually slightly smaller than or

FAMILY PLEUROBRANCHAEIDAE 3

equal to the foot in size, but sometimes it is larger. The relative proportions of

the mantle to the foot were used for generic separation but they are largely de-

pendent upon the degree of contraction during preservation. Anteriorly the no-

tum is separated from the oral veil by a slight fold. The notum may be smooth

or rough, or with bosses or tubercles, which may be altered during preservation.

The right side of the notum projects over the gill, forming an exhalant

siphon posteriorly. On the left side the notum sometimes leaves the upper side

of the foot free. Posteriorly it merges gradually with the upper side of the foot.

In Pleurobranchaea brockii the notum and foot remain separated posteriorly.

The oral veil is broad. Its anterior border bears one or two rows of simple

or branched papillae, which may disappear with fixation. The sides of the veil

form the oral tentacles, which are split posteriorly and inwardly rolled. The

blunt, rolled rhinophores are situated in the angles between the tentacles and

lateral notal borders and are well separated from each other.

Between the right rhinophore and the gill are the genital apertures, sur-

rounded by a circular fold. This fold often has one or two flaps that are fre-

quently protruded. The flaps vary intraspecifically. Behind these apertures lies

the prebranchial gland, Bourne’s gland, called the organ of Bojanus by Moquin-

Tandon (1870: 20). It opens on a small papilla.

The gill occupies the middle of the right side below the border of the no-

tum. It has a variable number of pinnae on both sides, which increases with age.

The gill rachis is smooth or bears two rows of papillae, which may disappear

with preservation. Part of the underside of the gill is adnate to the body wall.

The anal opening is situated between the gill and notum, while the nephropore

is ventral to the gill.

The anterior border of the foot is transversely grooved. The hind end is

rounded and often bears a soft spur or horn on the dorsal side. The presence or

absence of the spur (Figs 6A, 13A) varies intraspecifically. The pedal (metapodi-

al) gland on the ventral side of the foot is not always visible. Macnae (1962:

168-169) found this gland only in sexually active individuals, although its ab-

sence may be a result of poor preservation.

The mouth opening is often extended by the everted buccal cavity and pro-

truded pharynx. The pharynx has a vertical entrance to the digestive tract and is

flanked by the jaw plates. The anterior border of the jaws may vary in shape

(Figs 6G, 19C) even in the same species (Marcus, Ev. & Marcus 1966, fig. 35).

The platelets comprising the jaws are polygonal columns with a flat surface. The

ratio of length to breadth of the platelets varies intraspecifically and may vary

with position in a single individual (Fig. 10C-F). We agree with O’Donoghue

(1929: 58) that the jaw platelets cannot be utilized for specific separation.

The radula has numerous transverse rows of uncinate, uni- or bicuspid

teeth. A rachidian tooth is rarely present. The teeth are largest in the middle of

the half-row. The presence or absence of a secondary denticle on the teeth is

variable. The number of unicuspid teeth on the outer edge of the radula also

varies. In some specimens of P. tarda there are up to twenty unicuspid teeth,

4 ANNALS OF THE SOUTH AFRICAN MUSEUM

while in other individuals they are entirely absent. We believe that the rudi-

mentary secondary cusp on all teeth in P. californica is a useful subgeneric

character. In specimens where the radular rachis is more or less complete, the

first lateral tooth varies in size in different rows. It gradually increases in size

from older teeth to newer and ranges from 0,105 to 0,250 mm. As additional

teeth are added to the radula from the inner side, the size of the inner lateral

again decreases. Malformations of teeth are not rare. Most commonly two

adjacent teeth are fused within a transverse row. On one occasion two entire

transverse rows of abnormal teeth were observed (Fig. 13B).

The stomach often contains the remains of food, demonstrating the vari-

ability of diet within the Pleurobranchaeidae. In the specimens examined, prey

included the bryozoan Cellaria sp., polychaetes up to 15 mm long, an amphipod,

a small prosobranch shell, a specimen and a pharynx of a pleurobranchid, a

gymnosomatid radula, an aeolid 2,5 mm long and a complete pharynx with jaw

plates and radula, a 20 mm branch of a crinoid, a spine of the sea urchin Eucida-

ris tubulinoides (identified by F. M. Bayer, U.S. National Museum) and a 6 mm

long octopod. A 65 mm specimen of Pleurobranchaea brockii contained four

specimens of Philine aperta in its stomach, the largest of which was 18 mm in

length. Eales (1937) found fourteen specimens (10-17 mm long) of Pleurobran-

chaea tarda in the stomach of Pleurobranchella nicobarica.

Bergh (1897: 2) indicated that a blood gland is situated dorsal to the heart

in the Pleurobranchacea, not over the central nervous system as in the Dorida-

cea.

Bergh’s (1897: 25-26, pl. 4 (figs 1-7)) description and figures of the cuticu-

lar penial tube of P. meckelii differ from Vayssiére’s (1901: 40-41, fig. 228). As

this is the evaginable part of the male duct, we consider it as the penis proper.

Bergh’s (1897: 34-37) terms for the reproductive ducts differ somewhat from

ours. Vayssiere (1901: 39-40, pls 4-6) described the male duct of P. meckelii.

He called Bergh’s term ‘penial sheath’ the penial sac (his fig. 224) ard applied

the term ‘penis’ rather than ‘glans penis’ to the small muscular cone at the open-

ing of the efferent duct into the atrium (his figs 225-226). He did not mention

nor figure any cuticularized penis inside the efferent duct, except for a trans-

verse section (fig. 228). Macnae (1962) roughly figured the penis of four pleuro-

branchid species. MacFarland (1966: 99, pl. 17 (figs 1-7)) described and figured

the ‘glans penis’ of Pleurobranchaea californica and compared it with Bergh’s

description of P. meckelii.

The inner genital organs dissected from the intestine and digestive gland,

but still connected with the body wall, can best be studied by slightly staining

them in carmine and sketching them before clarification to see the cuticular

penis, if present. In laying the organs flat (Vayssiére 1901: 38) for balsam prepa-

rations they frequently stick together or tear, so that the course of the ducts is

no longer recognizable.

From the ovotestis, in which the follicles contain either sperm or oocytes

(Mazzarelli 1891: 234), a winding hermaphroditic duct emerges. It widens as an

FAMILY PLEUROBRANCHAEIDAE >)

elongate ampulla. Shortly after it narrows again, it divides into the oviduct and

male duct.

The ciliated oviduct widens, forming some pouches full of sperm, the sper-

matocyst (Fig. 3B, y). The following long, coiled portion is also ciliated. The

oviduct enters the vagina near the stalk of the vesicular spermatheca. The albu-

men and mucous glands open into the vagina.

The male duct generally enters the centre of a round prostatic gland com-

posed of many narrow tubules. It emerges from the gland on the opposite side

and enters the penial sac, when present, usually near its fundus and often close

to the retractor muscle. In Euselenops the prostatic cells are contained with the

efferent duct.

The penial sac is narrow along the loop of the efferent duct in P. tarda and

slightly wider in P. maculata, P. bubala, and P. californica, and widest in P.

meckelii and P. augusta. In one specimen of P. notmec and in P. bonnieae and

P. inconspicua, it is tight around the many windings of the efferent duct (Fig.

18F). The number of crowded windings of the efferent duct in the penial sac in

P. inconspicua, P. bonnieae and P. notmec could not always be accurately deter-

mined. The penis is not known in detail for P. maculata. The penis does not

have a cuticle in P. agassizii, P. brockii, P. bonnieae, and P. vayssierei. It is cov-

ered with papillae in Euselenops luniceps, and with cuticular hooks in Pleuro-

branchella nicobarica (Fig. 24D) and Koonsia obesa. There is an external cuticle

in Pleurobranchaea notmec (Fig. 18C—E), and an internal cuticular cylindrical

stylet of different transverse section (Fig. 1A—-H) in Pleurobranchaea meckeiii,

P. tarda, P. inconspicua, P. californica, and P. bubala.

The spermatocyst is often a simple widening, or its epithelium forms

pouches. In P. brockii it is generally a spiral caecum of three to five whorls, but

in one specimen (SAM—A29866) it consisted of epithelial pouches.

The spermatheca is small and not transparent, or wide and thin-walled. In

P. bubala there may be a larger ental vesicle with a smaller ectal one on its top

(Fig. 16G).

Mazzarelli (1891: 233-238, figs 1-3) gave an excellent description of the re-

productive apparatus of P. meckelii, but seemed to have confused the prostate

and albumen glands. He described the cuticular tube in the efferent duct,

though he called it a filament. By dissecting mating specimens he found that the

cuticular tube is projected from the everted penial sheath into the widenings of

the inner oviduct of the partner, the spermatheca and spermatocyst.

KEY TO THE NOTASPIDEAN GENERA

(Pe Shiclifextental Serer eee eee a ie nah s ae oa ko Phe de dee ee ee Z

SS ONICHIMINGeKMalKOtmADSCHt ame can acetate melee orn ced gts ss ene eens ne Som ses 5

Pee Shelliparthycovercdibyamantlowe ws... setae ce sce shee eee che dees tee Gymnotoplax

eo nellinotcovercdibyamantic(@Wmbraculacea). ¢..gss4s-eec8s0ss eh ode oases bee oe ae 3

3. Gill from left rhinophore along front and most of right side............... Umbraculum

Se OlllLonlyonypanvomunemiCNNSIdCre ta abe tsa te om bene gerd ee he ees eee tes de mee +

JNA GAM LOO WPESeMt teat sioe ts oe aia kiko awit Peat bd ee coe ee wo de Tylodina

INO brachicianecoothiearewrag eon ices es Us te AES waned NE Ml aoe sbeaais & dake ealie odechases Roya

6 ANNALS OF THE SOUTH AFRICAN MUSEUM

5 Shell internal; rhinophores anteromedial, posterior to veil; jaw elements flat posteriorly,

pointedianteriorly (Pleurobranchidae) 42455 7 sees ae ee ee 6

Shell absent, rhinophores lateral, between veil and mantle border; jaw elements more or

less high polyconal'column'(Pleurobranchaeidac) a sees ee eee 11

6) Oviduet enters'albumen)eland|(Berthellinac)-ee ss ee ee 7

— ‘Oviduct unites with vagina (Rleurobranchinae)ie ao. 4 .4oe eae ae ee eee 9

7 ~Yeeth lamellitorm . 5 2.0. + oye esas Fo ha oe Spe ee ee On 8

—. Teeth hook shaped...) 0 sce oe nuh aoe te ane Neo ee ae Berthella

8 Teeth with one secondary cusp; prostatic gland cells in male duct........... Pleurehdera

——) Neethidenticulate prostate presenter ee a eee eee eee Berthellina

9° “Prostate presemt: 5 i 2. chine « artaeiaye a Som cece OS Sucre tise oe ne 10

=" Prostatic, sland cells;in: male: duct; 23-447 so ne ea a eee Susania

LO} Shelllarge-ibig fapsvaround!genitalvapentunesi ese en eee Oscanius

— Shell small, no big flaps around genital apertures...................... Pleurobranchus

11 ‘Notum half the length of f00t+-5 0. 25.3 dee en oe nee on eon Cr 12

—* Notum larger than footor equal... 2c pansies oe ee oo eee 13

12S Boldionhindtendloknotumspenisipapillateaeen eee eee Euselenops

=" Spuron hind endo toot] penisswith hoOksiee naa ne eee eee Koonsia

13) Radular teéeth:unicuspide © 5.235 ee eee eee re eee Pleurobranchella

— | Radular'teethibicuspid 2.35506 San ee ae ee ee 14

14 Secondary cusp of tooth well developed............ Pleurobranchaea (Pleurobranchaea)

= Se candahy Cusp iuGiINentahy nal are an eae ae Pleurobranchaea (Macfarlandaea)

SPECIES OF THE FAMILY PLEUROBRANCHAEIDAE TREATED IN THE PRESENT PAPER

Pleurobranchaea meckelii Leue, 1813; Mediterranean to Gulf of Guinea

. maculata (Quoy & Gaimard, 1832); Australia, New Zealand, Japan

. tarda Verrill, 1882: 546; western and south-eastern Atlantic

. agassizii Bergh, 1897; western Atlantic

brockii Bergh, 1897: 41; Japan to South Africa

. Inconspicua Bergh, 1897: 49; western Atlantic, Mediterranean, Israel

confusa sp. nov. pro parte Koonsia obesa, Verrill; north-western Atlantic

augusta sp. nov.; west Africa

bonnieae sp. nov.; Florida

bubala sp. nov.; Sow Africa

notmec sp. nov.; Israel

. vayssierei sp. nov.; Algiers

. (Macfarlandaea) subgen. nov.

. (M.) californica MacFarland, 1966: 94; California

Busclenons luniceps (Cuvier, 1817); Japan to South Africa

Koonsia obesa Verrill, 1882: 545; 1884, pl. 28 (fig. 7); western Atlantic

Pleurobranchella nicobarica Thiele, 1925: 283; Indian Ocean, South Africa

Gigantonotum album Lin Guangyu & Tchang Si, 1965; China

nel ssh nel as} ae) ae) ns} os} Fe) Sel Se) a5) Mel

LIST OF SYNONYMS

Pleurobranchaea meckelii var. occidentalis Bergh, 1897: 28; syn. nov. of P. tarda

Pleurobranchaea hedgpethi Abbott, 1952; syn. nov. of P. inconspicua

Oscaniopsis semperi Bergh, 1897: 54; syn. of Euselenops luniceps

Oscaniopsis compta Bergh, 1897: 58; syn. of Euselenops luniceps

Oscaniopsis amboinei Vayssiére, 1901: 15; syn. of Euselonops luniceps

FAMILY PLEUROBRANCHAEIDAE 7

Pleurobranchaea capensis, Vayssiére, 1901: 46; syn. nov. of P. tarda

Pleurobranchoides gilchristi O'Donoghue, 1929: 62; syn. nov. of Pleurobranchel-

la nicobarica Thiele

Pleurobranchaea hamwa Marcus & Marcus, 1955: 21; syn. of P. inconspicua

Pleurobranchaea gemini Macnae, 1962: 178; syn. nov. of P. brockii

Pleurobranchaea gela Marcus & Marcus, 1966: 174; syn. nov. of P. inconspicua

Pleurobranchaea occidentalis Bergh, 1897: 28; syn. nov. of P. tarda

Pleurobranchaea novaezealandiae Cheeseman, 1878: 378; syn. of P. maculata

INSUFFICIENTLY DESCRIBED SPECIES NOT TREATED HERE

Pleurobranchillus morosus Bergh, 1892: 28

Pleurobranchaea morula Bergh, 1905: 48

Pleurobranchaea melanopus Bergh, 1907: 33

Oscaniopsis pleurobranchaeana Bergh, 1907: 35

Pleurobranchaea algoensis Thiele, 1925: 282

Pleurobranchaea japonica Thiele, 1925: 283

Pleurobranchaea dorsalis Allan, 1933: 445

Classification, synonymy, and geographic distribution of the insufficiently

described species remain doubtful, .particularly as two or more well-

distinguished species may be sympatric.

[Sl Slelealalalwls|wlele

KEY TO THE SPECIES OF THE FAMILY PLEUROBRANCHAEIDAE

INotumbhalitthescnethtotthetoote asi aren oe eine mee esate eG dete be eee be ae Z

INosumblancemthanioovonecqualieacsa 4 a. eos ate eis oe nee ted eee oe et ew eae | 3

Roldvonthindrendlofnotums penis papillate aye. sae aaa ee Euselenops luniceps

SPUmon mincdienadoltoot penis withiiOOks) 4.4.46 eae. es lo. -o- Koonsia obesa

aduilarteethtumicuspidh.. sea ae. aoe cae oee cs ener tae mee Pleurobranchella nicobarica

Naa Ul arte e Chiat Cus Picline wanes cer ree an hs A wee Poe a Ma ee LPS ches, geome 4

Secondary cusp of tooth well developed ...... . Pleurobranchaea (Pleurobranchaea) .... 5

Secondary cusp rudimentary.............. Pleurobranchaea (Macfarlandaea) californica

REMISISOLMECULIC ORD SC It eran Were ware nie hed ae ie EGON Le ie sed oy bade a 6

ISHS CUCM Sis a ois dee Gea: Bet cl oL Orly te En ORNS yee to once MEER IT

WATE S NOME <5 3 glee, bo 68 bd acd Ao Oo Hiro Mid citer HIG isi cea Gane ees arene ne ae ete besten 7

VEU MOVIE 5 6 dio: doo. ove Odo. od G15, Gro e URG eS aOR ark res ls otc a rae ea 10

IN@® [DEMME GAC.ou; a wig bbe Blac sig bots ee Magrerore Bromo See one toate RR ann P. agassizii

enialisa cd ev elope din: qymermrnen cro eee ie MRSS Nas dss Ales ca tuayh a Heh bee tees 8

RE MIAlESA CH ARKOWareite aay ware te ae eM yl a cade SE Ss P. maculata

PSTN GAGS CUOITIBNE os.°5 Shoso Goto rciote os 8) 6'b polols ra)n)G Gees Gln S IO cope ne Rater earner afc et 9

NOOMOl PE MISIeMlAanged ian. wanes eee ens eas edie he ews Be eas P. bonnieae

IOOUOL penismot widen than following parte... ee s4.¢e5-2-62+ 0405+. 5: P. vayssierei

eMISAVehys ONG ran GepniChpmmrnr ase eee ite Fa Ge en aise pase bas eo ates P. brockii

JOINS GINO soe oles 6 oo RAUMLaOm 6 Gta Rinne ons CENCE Rae een eee ee P. confusa

memistarsligintlyscumvc GisOViCtene at. cma yee ee mee Ce ee eee cw lnt ovla Seated o's ween ofl 12

Renisa lone ube withaanyloopsin)pemialiSaci anys) 458555) ae neste eos s oe 13

ESMIASAC Manowar till ania. (4 ee oe yk ee sa kins cbs m vues LHe wera hare P. tarda

Reimialsac widencdrarOunquGuct euaesnecm cas lem.. aqee na cess hase cena P. bubala

\WRGANI EY SINOLEL’ <a io & cls ooo: accied Notice Sass nine eT ee ed et an Sone ere ares a ere 14

NYRI OTN D386, oto 6 dig orcs eho Bless ol Ghee cients noe tea ae ea P. meckelii

Penial cuticle with crest, less than ten loops in penial sac ................... P. augusta

Penial cuticle cylindrical, ovoid in transverse section, up to twenty loops in penial sac

P. inconspicua

8 ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarks

Sufficient characters were not obtained to include P. notmec sp. nov. in the

key, though in the discussion it is distinguished from similar species. Bergh’s

(1892: 27, 28) genus Pleurobranchillus for morosus and brockii was synonymized

to Koonsia by Pilsbry (1896: 221, 223) and by Abbott (1974: 349). Later Bergh

(1897: 27, 28) transferred morosus and brockii to Pleurobranchaea.

Thiele’s (1925: 283) Pleurobranchella and its synonym Pleurobranchoides

O’Donoghue, 1929, have a notum noticeably larger than the foot. This is excep-

tional in the family Pleurobranchaeidae.

Pleurobranchaea Leue, 1813

Pleurobranchidium Blainville, 1824.

Cyanogaster Blainville, 1825.

Pleurobranchillus Bergh, 1892.

Koonsia Verrill, 1882, partim.

Non Pleurobranchella Thiele, 1925.

Type species

Pleurobranchaea meckelii Leue, 1813.

Diagnosis

Pleurobranchaeidae with bicuspid radular teeth.

Pleurobranchaea (Pleurobranchaea) Leue, 1813

Type species

Pleurobranchaea meckelii Leue, 1813.

Diagnosis

Secondary cusp of radular teeth well developed.

Remarks

Thiele’s (1931: 419) definition is confused and the forms he attributed to the

subgenus have quite different characters.

Pleurobranchaea meckelii Leue, 1813

Figs 1A, H—-P, 2A—C

Pleurobranchaea meckelii Leue, 1813.

Pleurobranchidium meckelii Blainville, 1824.

Pleurobranchaea meckelii Leue, 1813. Mazarelli, 1891: 223. Pelseneer, 1894: 31, figs 86-99.

Bergh, 1897: 7, pls 1-4, pl. 7 (figs 16-18); 1899: 26; 1902: 228, pl. 9 (figs 1-3). Vayssiére,

1901: 42, figs 205-231.

Non P. meckelii var. occidentalis Bergh, 1897: 28 (= P. tarda).

FAMILY PLEUROBRANCHAEIDAE 9

Fig. 1. A-H. Optical transverse sections of penial stylets. A, H. Pleuwrobranchaea meckelii

Leue, 1813. B. P. inconspicua Bergh, 1897. C. P. augusta sp. nov. D. P. bonnieae sp. nov.

E-G. P. notmec sp. nov. I-P. P. meckelii Leue, 1813. I-J. Jaw platelets from Algiers specimen

from Vayssiére’s collection. K-L. Teeth from Algiers specimen. M-—P. Jaw platelets from

Israel specimen.

10 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 2. A-C. Pleurobranchaea meckelii Leue, 1813. A. Penial sac. B. Optical section of

penial stylet. C. Diagram of reproductive organs. D. P. maculata (Quoy & Gaimard, 1832),

diagram of reproductive organs (after Vayssiére 1901, fig. 247).

Material

Museum National d’Histoire Naturelle, Paris: two specimens, Algiers, AI-

geria, 1900; five specimens, Mediterranean, Israel to northern Sinai. Zoology

Museum, Copenhagen: some dissected incomplete specimens. United States

National Museum, Washington, D.C.: P 239, one specimen, Gulf of Guinea

(04°56’N 05°00’E).

FAMILY PLEUROBRANCHAEIDAE it

Distribution

Mediterranean, Atlantic, Azores (Bergh, 1899); Cape Verde Islands (Vays-

siére, 1902).

Description

Length up to 100 mm. Radular teeth bicuspid, one to five outermost teeth

unicuspid. The efferent duct passes transversely through the retractor muscle. In

the strongly muscular penial sac it divides into penis proper and penis sheath.

The penis is an elastic cuticular stylet, 0,06—0,1 mm diameter, with a high crest

(Fig. 1A, H). It forms no more than six to ten loops and is up to 10,5 mm long.

Vayssiére’s (1901, fig. 223) figure of the structure of the penis, seen by

transparence, evidently refers to a young specimen.

Discussion

Bergh’s (1897) and Vayssiére’s (1901) descriptions and figures of P. mecke-

lii differ. Vayssiére evidently did not have homogeneous material. He indicated

a thin, transparent penial sac in P. meckelii, but for one of his samples an

Opaque, nacraeous, muscular one is present. He did not see the elastic cuticular

stylet that we found in his Algerian specimen (Fig. 2B) and in the Israeli speci-

men (Fig. 1A, H). The latter agrees perfectly with Bergh’s description and

figure (pl. 4, fig. 4) of the penial stylet, while the present specimen from Algiers

has a different transverse optical section.

Vayssiére (1901: 43) indicated that P. meckelii was the only species occur-

ring in the Mediterranean. Therefore, all subsequent findings of Pleurobran-

chaeidae from the region were identified as P. meckelii. However, the present

material from Turkey, Israel, and Algeria contains several other species. Even

among the three specimens received from the Museum National d’Histoire

Naturelle, of Vayssiere’s material collected in Algiers in 1890, there was one dif-

ferent from P. meckelii. It is called here P. vayssierei (see p. 38). The specimen

from the Gulf of Guinea extends the range of the species. Most of the synonyms

assigned to P. meckelii are insufficiently described. This is true of Sturany’s

(1904) description, and the record of specimens from the Red Sea is question-

able (Pruvot-Fol 1933: 91). P. meckelii var. occidentalis Bergh (1897: 28) is ac-

tually P. tarda.

Pleurobranchaea maculata (Quoy & Gaimard, 1832)

Figs 2D, 25E

Pleurobranchidium maculatum Quoy & Gaimard, 1832: 301-302, pl. 22 (figs 11-14).

Pleurobranchaea maculata (Quoy & Gaimard, 1832) Vayssiére, 1901: 49-56, figs 238-247.

Pleurobranchaea novaezealandiae Cheeseman, 1878: 378. Willan, 1983: 254, figs 57-70.

12 ANNALS OF THE SOUTH AFRICAN MUSEUM

Distribution

Australia, China, Japan, New Zealand, Sri Lanka.

Description

The length is up to about 50 mm.

The radular formula is 40—49 x 80.0.80; ten to fifteen of the outermost teeth

lack a secondary denticle.

The reproductive organs described by Vayssiére (1901) differ from those of

the other now known species by the long vagina ending in a globular caecum to

which the spermatheca is connected (Fig. 2D). The efferent duct winding up and

down the retractor muscle in the penial sac is similar to that of P. tarda (Fig.

5B), but it evidently does not have a stylet.

Discussion

Vayssiére (1901: 51) studied specimens of Quoy & Gaimard’s (1832) orig-

inal material and found that Bergh’s (1898b: 429, pl. 29, figs 2-9) animals from

Juan Fernandez were not P. maculata. The same must evidently hold for Odh-

ner’s (1921: 224) undescribed specimens from Masatierra.

The reproductive organs of a young specimen of P. tarda were somewhat

similar to those of P. maculata in Vayssiére’s figure, but the spermatocyst and

spermatheca are quite different.

The synonymizing of insufficiently described species only because they also

come from the Indo-Pacific region (Thompson 1970) cannot be maintained. P.

melanopus Bergh (1907: 33), P. japonica Thiele (1925: 283), and P. dorsalis Al-

lan (1933: 445) are possibly synonyms, but P. tarda (Fig. 5) is quite distinct from

P. maculata. Pleurobranchoides gilchristi O’ Donoghue (1929: 62), a synonym of

Pleurobranchella nicobarica Thiele (1925: 283), is a species with unicuspid teeth

and is also very different from P. maculata.

The male organ of Vayssiére’s (1901: 49-56, figs 238-247) P. maculata

(Quoy & Gaimard, 1832) from Australia is similar to that of Bergh’s (1897, pl.

7, fig. 14) P. tarda, but it does not have a stylet.

Pleurobranchaea confusa sp. nov.

Fig. 3

Koonsia obesa Verrill, 1882: 545-546, partim. Verrill, 1884: pl. 28 (fig. 7). Verrill, 1885, partim.

Pleurobranchaea obesa Bergh, 1897, non Verrill: 30-33, pl. 7 (figs 19-21). Vayssiére, 1901, non

Verrill.

Type material

Holotype: United States National Museum 784657, off Delaware Bay

(38°35'N 73°13'W), 400 m, 10 October 1881. This specimen was previously a

paratype of Koonsia obesa.

FAMILY PLEUROBRANCHAEIDAE 13

ES aa ares

= =

Se Sener fy

SOLS SS aaa

/}»

Fig. 3. | Pleurobranchaea confusa sp. nov. A. Ventral view of preserved specimen (paratype

of Koonsia obesa). B. Diagram of reproductive organs. C. Optical section of efferent duct in

penis. D. Tip of penis.

14 ANNALS OF THE SOUTH AFRICAN MUSEUM

Further distribution

? Off Martha’s Vineyard, Massachusetts.

Etymology

The epithet confusa (from Latin: confusio) refers to the combination of two

distinct species in the original description of Koonsia obesa.

Description

The present contracted animal is 4,5 by 4,0 cm and has a smooth, swollen

notum extending far out over the foot (Fig. 3A). The oral veil has a single row

of papillae. The tentacles and rhinophores are smooth. The triangular foot is

narrower and shorter than the notum. It has a broad anterior border, a dorsal

spur, and a ventral foot gland (Fig. 3A). The gill has about twenty-six pinnae

and a smooth rachis.

The denticles on the jaw platelets are irregular in presence and in size;

there are two to twelve denticles per platelet. The radular formula is

34 x 90.0.90. Approximately fifteen of the outermost teeth of each row are uni-

cuspid. There is no rachidian tooth.

The male duct enters the prostate and, after leaving it, winds to enter the

wide penial sac. Inside the sac it forms a wide sheath and a strongly muscular

penis with two tiny cuticular bulbs at its tip (Fig. 3D). The oviduct has some epi-

thelial pouches, the spermatocyst. The oviduct widens and receives the large

spermatheca. The wide vagina continues, joining with the female gland mass,

and opens through the female aperture (Fig. 3B).

Discussion

This description is based on a paratype of Koonsia obesa from Verrill’s col-

lection and corresponds to the paratype studied by Bergh (1897: 30-33, pl. 7

(figs 19-21)). It differs from Verrill’s (1884, 1885) figures by the swollen and

overhanging mantle, elongate tail and the absence of ‘small hooks on the verge’.

Bergh was quite right to transfer this material to Pleurobranchaea. However,

specimens of the type species of Koonsia, K. obesa Verrill, 1882, may be found

again. The name Pleurobranchaea confusa sp. nov. is proposed for the paratype

of Koonsia obesa.

Pleurobranchaea tarda Verrill, 1880

Figs 4-5

Pleurobranchaea tarda Verrill, 1880: 398; 1882: 546, pl. 58, (fig. 26). Bergh, 1897: 33, pl. 6 (figs

28-31), pl. 7 (figs 1-15). Vayssiére, 1901: 57, figs 248-251.

Pleurobranchaea tarda var. Bergh, 1897: 39, pl. 7 (figs 21-27).

Pleurobranchaea meckelii var. occidentalis Bergh, 1897: 28. Vayssiére, 1901: 46.

Pleurobranchaea occidentalis Bergh, 1897. Marcus & Marcus 1967a: 45.

Pleurobranchaea capensis Vayssiére, 1900. Vayssiére, 1901: 46, pl. 4 (figs 232-237). O’Dono-

ghue, 1929: 48, figs 52-57. White, 1955: 173, fig. 6.

Pleurobranchaea maculata Thompson, 1970: 192, partim., non Quoy & Gaimard.

FAMILY PLEUROBRANCHAEIDAE 15

Fig. 4. Pleurobranchaea tarda Verrill, 1882, living animal.

Material

About 120 specimens in the U.S. National Museum and South African Mu-

seum; western Atlantic, from 31° to 5°N; eastern Atlantic from Ghana and the

Cape of Good Hope and False Bay; intertidal to 1 450 m.

Further distribution

From Martha’s Vineyard to south of Cuba; Angola and Kabinda; Agulhas

Bank (35°10’S 23’E), 500 m.

Description

The present animals preserved, are from 7,5 to 52 mm long. The mantle is

generally smooth and about the same size as the foot. On its right side the bor-

der often forms an exhalant siphon over the anal opening and the tip of the gill.

The tip of the foot often extends posteriorly (Fig. 5C). In some specimens it

bears a short spur. The metapodial gland on the end of the sole is rarely devel-

oped. The veil, tentacles and rhinophores are typical for the genus. The tuber-

cles of the veil are sometimes preserved. The proboscis is often protracted, and

the anterior border of the jaw plates are as variable as in P. gela (Marcus, Ev. &

Marcus 1966, fig. 35).

ANNALS OF THE SOUTH AFRICAN MUSEUM

‘yoodse JemusA “q ‘JAIS Jo weg -“q ‘wodse jesiog ‘dD

‘uouttoods onuepy U19}SeM JO SUPSIO DAT]ONpOIdaY ‘q “UsUIDOds ULLIOSIN JO suUeSIO dATIONPOIddY “VY “ZBI ‘[[IOA vpsv) vavyouvégosnafg °¢ “314

SD 3

tasers

a ates,

AN O42TA

FOILS

CLT

CY)

ay NM

FAMILY PLEUROBRANCHAEIDAE 17

The gill is adnate for about two-thirds of its length. The anus lies dorsally,

just in front of the gill membrane. The number of pinnules of the gill varies from

twelve to thirty pairs. They alternate and often begin with a tubercle, so that the

rachis appears tuberculate. In some animals the gill is regenerating. The genital

apertures lie in the middle of the body or slightly anteriorly. According to their

state of contraction or eversion they are flat, small and round, with or without a

flap. The male atrium is partly everted and the tip of the cuticular penis is pro-

jected in about twenty of the present specimens (Figs 5C, E). In an animal of

10 mm the tip of the cuticular penis was well developed and stuck out of the

male aperture.

The radular teeth are bicuspid except for about six of the outermost ones.

From the ampulla (Fig. SA—B) the thin common genital duct (0), divides

into male (e) and female (f) ducts. The latter recurves backward along the com-

mon duct. It is much wider initially, forming a ciliated serial spermatocyst. Vays-

siére (1901: 49) thought this was a caecum of the common duct, from which the

efferent duct went out. Farther outward the female duct, which functions as

both oviduct and insemination duct, is rather wide and has a high ciliated epithe-

lium. The narrow efferent duct is surrounded by a ball of fine branching prostat-

ic gland tubules (q). As it leaves these tubules, it widens and loops around the

male atrium, entering the narrow penial sac (s) and traversing the retractor mus-

cle (r) for up to 7 mm. It then turns outward and reaches the fundus of the atri-

um (a). The length of the efferent duct varies considerably along the retractor

muscle and within the atrium. If the penial sac remains intact during prepara-

tion, the aspect of the duct is as in Bergh’s (1897: pl. 7 fig. 14) figure. When it is

torn, it appears like Vayssiére’s (1901, fig. 249) figure. In its descending course

the efferent duct is separated into a sheath (h) and the cuticular penis proper

(p). This is a stylet of 0,12 mm diameter at its base, narrowing to 0,08 mm in di-

ameter. Its length varies from 1,9 to 6 mm. The chitinous penis enters the atri-

um through a small papilla. The penial cuticle is rough on the outer side (Fig.

5D). Vayssiére’s (1901, fig. 25D) figure of the tip of the penis corresponds to our

observations, but we cannot confirm his interpretation of its muscular nature.

The oviduct begins with a winding spermatocyst and continues to widen

slowly to the stalk of the spermatheca (t). The vagina (v) is short and wide and

receives the outlets of the albumen and mucous glands. The atrium is generally

long and narrow when imverted, or short and wide when partially everted. Its

epithelium contains transverse bands of glandular cells.

Discussion

Both Bergh and Vayssiére studied specimens from Verrill’s original ma-

terial, but neither of them observed the cuticular stylet. They figured only the

tip of the penis (Bergh 1897, pl. 7 (fig. 27); Vayssiére 1901, fig. 250). Vayssiere

considered the penis of his specimens (30-40 mm alive, 14-16 mm preserved) in-

completely developed. In the present material a preserved animal 7,5 mm in

length had a well-developed cuticular penial stylet. The inward loop of the effer-

18 ANNALS OF THE SOUTH AFRICAN MUSEUM

ent duct in Bergh’s (1897, pl. 7 (fig. 14)) figure is held together by the penial sac,

while in Vayssiére’s figure it is loose.

Bergh (1897: 28-30, figs 16-18) described Pleurobranchaea meckelii var. oc-

cidentalis from Martinique. Marcus, Ev. & Marcus (1967a: 45-48, fig. 5SA—-C)

identified specimens from Florida with Bergh’s form and raised this to species

level due to the different penial sac. Marcus & Marcus figured the penis in its

sheath (their fig. 55C). Comparing this and the present animals with P. tarda in

Vayssiére’s figure 249, the authors did not find any difference and consider

P. occidentalis as a synonym of P. tarda.

Pleurobranchaea capensis was described from two specimens from the Cape

of Good Hope collected in 1829. Vayssiére (1901: 46-49, figs 232-237) described

the inward loop of the efferent duct as being similar to that of P. tarda. How-

ever, it did not have a loop around the atrium (Fig. 5A).

Bergh (1907: 30) identified twenty-seven of his South African specimens as

P. capensis Vayssiére, with the ‘vas deferens much shorter than in P. meckelit’.

One specimen Bergh (1907: 32, pl. 11 (figs 1-8)) called P. capensis var., because

it had a ‘much longer vas deferens than represented by Vayssiére (fig. 232)’.

The present ample material showed a high variability of the length of the male

duct. The extremes corresponded to those described for P. tarda and P. capen-

sis. Hence P. capensis Vayssiére must be considered as a junior synonym of

P. tarda Verrill.

Pleurobranchaea brockii Bergh, 1897

Figs 6-8

Pleurobranchillus brockii Bergh, 1892: 28, nom. nud.

Pleurobranchaea brockii Bergh, 1897: 41-46, pl. 4 (figs 8-17). Vayssiére 1901: 62-69, figs

255-260. ? Farran, 1905: 355, pl. 5 (figs 24-28). White, 1948: 20. Macnae, 1962: 178. Lin

Guangyu & Tchang Si, 1965: 275, pl. 1 (fig. 3).

Pleurobranchaea brockii var. Bergh, 1897: 46-47, pl. 7 (figs 33-38), pl. 8 (fig. 1).

Pleurobranchaea gemini Macnae, 1962: 178, fig. 7a—c.

Material

South African Museum: SAM-—A29866, two specimens, no locality; SAM—

A30078, one specimen, no locality; holotype P. gemini, SAM—A35234, Inhaca

Island, collected by W. Macnae, 18 July 1958; paratype P. gemini, SAM-—

A35236, Inhaca, collected by W. Macnae, 19 July 1958; SAM—A35237, two

specimens and egg mass, Inhaca Island, collected by W. Macnae, December

1967.

Distribution

Japan to South Africa.

Description

The preserved specimens are 38-110 mm long. The notum is smooth or ma-

millate. The colour, when preserved, is a net of black pigment around more or

FAMILY PLEUROBRANCHAEIDAE 19

wee DPR ta sce sas ee

ROSCA het ea Per

SSO

“

—

Ase)

oe veleey

Fig. 6. | Pleurobranchaea brockii Bergh, 1897. A. Dorsal view of preserved specimen.

B. Ventral view. C. Extruded penis. D. Tentacle and rhinophore. E. Gonopore. F. Trans-

verse section of vagina, scale 0,5 mm. G. Anterior of pharynx with borders of jaws.

20 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 7. Pleurobranchaea brockii Bergh, 1897. A. Diagram of reproductive organs of speci-

men SAM-A30078. B.Spermatocyst. C. Tip of contracted penis. D. Female organs.

FAMILY PLEUROBRANCHAEIDAE pa

Fig. 8. Pleurobranchaea brockii Bergh, 1897, paratype of P. gemini. A. Dorsal view of pre-

served specimen. B-C. Reproductive organs. D. Gonopore with a partially everted atrium.

E. Everted male and female pores.

eps ANNALS OF THE SOUTH AFRICAN MUSEUM

less regular light circles or big blotches, which contain mamillae. Pigment is also

present on the upper side of the foot. The notal border is free around its entire

surface. There are sixteen papillae on the border of the oral veil arranged in two

rows. The anterior side of the tentacles is thickly papillate in the South African

Museum specimens (Fig. 6A, D). In those from Inhaca there are fewer tubercles

(Fig. 8A). The rhinophores are set in deep sinuses between the tentacles and no-

tum. The foot projects further posteriorly in the specimens from Inhaca (Fig.

8A), but this may be an artifact of preservation. A spur and a foot gland are

present or absent. The gill rachis is narrow; the base of the 30-40 pairs of pinnae

may bear a knob.

The jaw plates have a straight anterior border (Fig. 6G). Their denticles are

present or absent and are irregular in number and size. The radular formula

is 40-48 x 65—-90.0.65-90. The teeth are bicuspid but the outermost laterals

are sometimes unicuspid. Vayssiére (1901, fig. 260) mentioned and figured a

rachidian tooth, but said it was atrophied and often lost or absent. It seems to

be the fused first laterals. The radula of the holotype specimen of P. gemini 1s

missing. Those of the paratypes are, like that of P. brockii, without a rachidian

tooth.

The male duct passes through the prostate (Fig. 7A) and enters the narrow

penial sac (s). There it forms a strong muscular, non-cuticularized penis (p). The

retractor muscle inserts on the middle of the dorsum and traverses the sac to the

atrium. In a contracted state the penis is curled (Fig. 8C). When extruded it is

up to 4 cm long (Fig. 6C). The shape of the atrium could not be determined.

When everted it is a wide and very thin sac (Fig. 8D), and when completely ex-

truded the male and female pores open on its innermost stronger part (Fig. 8E).

The gonopore is surrounded by a circular fold with some flaps.

The oviduct, after separating from the common duct, has a swollen part

(Bergh 1897, pl. 4 (fig. 8)), that sometimes forms a small caecum lying at the be-

ginning of the spermatocyst (Fig. 7A). In most specimens the latter forms a

longer caecum with three to five spirals, but rarely it is only a widening with epi-

thelial pouches in the duct. The duct narrows into several whorls and then

widens suddenly as the vagina. The vagina is long (Bergh 1897: 46) and its epi-

thelium is divided into many folds (Fig. 6F). Farther outwards the spermathecal

pouch opens into the vagina. The vagina joins the albumen and mucous glands.

It often has an ectal widening on one side.

The holotype of P. gemini had previously been opened only for the radula.

The female organs were intact (Macnae 1962, fig. 7c). The male organ was

examined for comparison. It corresponds to Bergh’s (1897) figure. The spawn is

a tall convoluted ribbon 13 mm wide, like that of P. californica.

Discussion

The descriptions of P. brockii and P. gemini are similar, and Macnae’s dis-

tinctive characters are not convincing: P. brockii ‘is characterized by a mamelo-

nated mantle’ while the ‘smooth, not mamillated surface’ of P. gemini is not

FAMILY PLEUROBRANCHAEIDAE 23

mentioned in the description, only in the comparison. In some of Macnae’s

paratypes which we observed the mantle was smooth, in others mamillate.

In all available specimens the sixteen velar papillae are set in two rows. In

the South African Museum specimens SAM—A29866 and SAM-—A30078, the

tentacles are thickly covered with papillae (Fig. 6D), while in Macnae’s animals

there are only a few. This merely represents intraspecific variation. Bergh found

no trace of rachidian teeth in the radula of P. brockii. In Macnae’s (1962: 178)

specimen of P. brockii they were ‘very deciduous and all had fallen off. . . and

were found in the debris at the bottom of the watch glass’. In P. gemini they

were not mentioned. Macnae’s figure shows very long primary cusps and the sec-

ondary cusp inserting near the base, but he (1962: 180) noted that they were

‘without the basal prominence on the inner edge’. The teeth of the paratypes of

P. gemini that were examined are like those of P. brockii. The radulae of ten

specimens were studied and a basal prominence was found in all specimens. It is

sometimes not visible when the teeth are tilted and this fact may explain why

Macnae did not notice the prominence in the holotype.

The ‘long and worm-like penis’ of P. gemini described by Macnae (1962:

180) was not figured. Bergh (1897, pl. 4 (fig. 8)) and Vayssiére (1901, fig. 256)

illustrated the large, soft, non-cuticular penis of P. brockii. A second caecum on

the outer oviduct of P. gemini, to which the spermatheca is attached (Fig. 8B),

is not recognizable in Bergh’s figure of P. brockii nor in the dissected specimens

from Inhaca. Based on the variability observed in Macnae’s type material and

the similarity of it to P. brockii, the two species are thought to be synonymous.

Pleurobranchaea agassizii Bergh, 1897

Fig. 9

Pleurobranchaea agassizii Bergh, 1897: 48, pl. 7 (figs 28-32). Marcus & Marcus, 1967a: 49, 51,

fig. 56A—D.

Distribution

Straits of Florida; Great Bahama Bank, 262-620 m.

Description

Length up to 10 cm; radular formula 32 x 98.0.98. About seventeen of the

outermost teeth lack the secondary denticle. The efferent duct (Fig. 9E) does

not enter the penial sac but, where the retractor muscle is attached to it, it divi-

des into a sheath and a muscular penis without cuticle. (After Marcus, Ev. &

Marcus, 1967a).

Discussion

In Marcus & Marcus’s (1967a: 49-51) description there were two errors.

P. 51: the allosperm duct does not rise through the efferent duct but through the

vagina and insemination duct, which is the outer oviduct. In figure 56D the ovi-

24 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 9. Pleurobranchaea agassizii Bergh, 1897, Reproductive organs.

duct is shown entering the albumen gland; actually, it only passes near the fe-

male gland mass and continues to the vagina, where it receives the secretions of

the albumen and mucous glands.

Pleurobranchaea inconspicua Bergh, 1897

Figs 1B, 10-11

Pleurobranchaea inconspicua Bergh, 1897: 49, pl. 8 (figs 2-10).

Pleurobranchaea hedgpethi Abbott, 1952: 1, pl. 1 (figs 1-8). Marcus & Marcus, 1959: 253,

fig. 6. Marcus, Er., 1961: 141. Marcus & Marcus, 1967b: 200. Marcus & Marcus, 1969: 18.

Nijssen-Meyer, 1965: 143, figs 1-3.

Pleurobranchaea gela Marcus & Marcus, 1966: 174, figs 35-37. Marcus & Marcus, 1968: 1336.

Material

About twenty-five specimens from 37°N to 5°N, 54°W to 86°W, 27-310 m;

one specimen from Atlit, Israel.

Further distribution

From Cape Hatteras to Argentina; Texas; Ivory Coast to Nigeria.

Description

The preserved animals range from 11 to 55 mm in length and are in various

states of contraction. Often the buccal mass is everted around the large

FAMILY PLEUROBRANCHAEIDAE DS

Fig. 10. Pleurobranchaea inconspicua Bergh, 1897. A. Everted penis, scale 2,0 m

B. Jaws showing location of platelets in C-F. C-—F. Platelets from various locations. G. L

eral view of platelet. H. Penis of specimen from Israel.

26 ANNALS OF THE SOUTH AFRICAN MUSEUM

Ze Dr, 4 MLD q

Mpotr wey NN

HE) a

cen,

h AOE,

Fig. 11. | Pleurobranchaea inconspicua Bergh, 1897. A. Diagram of reproductive organs.

B. Root of penis and penis sheath.

projected pharynx. The anterior borders of the jaw plates are then visible, and

sometimes even the radula. Also the male atrium can be everted (Fig. 10A), and

the cuticular penial stylet may be observed. The notum is smooth or has some

bosses separated by pigment, which is sometimes retained in these furrows.

The jaw plates of a 15 mm long animal were 4,6 X 2 mm. In a 26 mm speci-

men they were 5 X 1,6 mm. They are colourless in small specimens and light

brown in the largest ones. The platelets are composed of columnar rodlets

whose surface is often hexagonal. Their length varies according to their position

in the jaw and with the size of the animal. The anterior border has a row of five

FAMILY PLEUROBRANCHAEIDAE Mg

to fifteen denticles and often some knobs on the surface. These both can be lost

in the anterior, oldest part of the jaw plate.

The radular formula is 32—40 x 55-68.0.55-68. The lateral teeth are bicus-

pid as in other species of Pleurobranchaea, largest in the middle of the half-row

and smaller towards the outer border. Up to twenty outer teeth lack a secondary

denticle.

The ovotestis and hermaphroditic duct are typical of the genus, as are the

male duct and prostate (Fig. 11A). The male duct is 0,13 mm in diameter. Be-

yond where the male duct enters the penial sac, it is surrounded by a sheath

(Fig. 11B). Its lumen is cuticularized and stiffens the duct so that it forms up to

twenty roundish loops (Fig. 10H). This is the eversible penis proper, which may

reach 14 mm in length when extended. It is ovoid in transverse view, and flat-

tened on one side. Its lumen is circular or higher than broad (Fig. 1B). Ectally it

gradually tapers to its tip. It is difficult to find the point where the penial sheath

is separated from the penis proper, owing to the many loops of the efferent

duct. Therefore it was not distinguished in the figures of the penial sac. The

sheath is about 0,34—0,40 mm in diameter at its outer part. It ends forming a

small papilla in the fundus of the male atrium. The papilla is up to 7 mm long

and 2 mm wide. The entire male organ reaches 20 mm in length. The atrium can

be everted for copulation (Fig. 10A) and the cuticular stylet projected. The

smallest animal with a cuticular penis was 11 mm long. In material from 40°S, up

to 20 mm long, specimens still did not have genital apertures (Marcus, Ev. &

Marcus 1969: 20).

Discussion

Pleurobranchaea inconspicua, of which Bergh had a single specimen from

Brazil (Sergipe), is identical in all its described characters with P. hedgpethi Ab-

bott, 1952. Hence it is considered as a senior synonym of P. hedgpethi. The in-

semination ducts of P. hedgpethi (Marcus, Ev. & Marcus 1959, fig. 6) and

P. gela (Marcus, Ev. & Marcus 1966, fig. 37) were figured differently. These dif-

ferences appear to be a result of their state of contraction. Marcus, Ev. &

Marcus (1967a: fig. 56) illustrated the insemination duct of P. hedgpethi. It has

two vesicles as described for P. gela. In the present collection both forms occur.

It is believed that P. gela is also a junior synonym of P. inconspicua.

Pleurobranchaea augusta sp. nov.

Figs 1, 12

Type material

Holotype: United States National Museum 809995, West Africa (17°02'S

11°40'E), 54 m.

Etymology

From the Latin augustus, meaning venerable.

28 ANNALS OF THE SOUTH AFRICAN MUSEUM

Os)

Fig. 12. Pleurobranchaea augusta sp. nov. A. Jaw platelets from middle of jaw, scale

40 um. B. The same, from front of jaw, scale 50 um. C. The same, from near border of jaw.

D. Preserved animal. E. Outermost teeth. F. Radular tooth from middle of half-row.

G. Female organs. H. Penial sac.

FAMILY PLEUROBRANCHAEIDAE 29

Description

The animal is 32 mm long, 20 mm broad, and 15 mm high, in the preserved

state (Fig. 12D). It is colourless. The smooth notum forms an exhalant siphon

over the tip of the gill. It is smaller than the foot. The foot has a tiny spur and a

weak metapodial gland. The oral veil, tentacles and rhinophores are contracted.

The gill has a smooth rachis with about twenty pinnae on either side. On the

base of each of the pinnae there is a knob. Four-fifths of the length of the gill

are adnate. The labial cuticle bears conical papillae. The platelets of the jaws

vary in shape according to their position on the jaw. The central ones (Fig. 12B)

have a long narrow surface with about ten denticles and some small tubercles

behind them. The marginal platelets appear roundish with less numerous and

shorter denticles (Fig. 12C). The radular formula is 40 x 70.0.70. The teeth are

bicuspid (Fig. 12F) except for one or two of the outermost ones (Fig. 12E).

Ectally from the ampulla the hermaphroditic duct divides into the male

efferent duct, and the oviduct (Fig. 12G). The former passes through the pros-

tate and enters the thin, roundish penial sac in its middle. The efferent duct

winds upward to the entrance of the retractor muscle which it penetrates (Fig.

12H). Later it divides, widening slightly, into the penial sheath and cuticular,

stiff penis proper. Together the penis and its sheath form about six loops. In the

present specimen the penis ends before the sheath opens into the atrium with a

small papilla.

The penial cuticle is cylindrical with equally thick walls. It forms a sharp

crest (Fig. 1C), which is visible in optical transverse section. This crest serves as

the distinctive character separating P. augusta from P. meckelii, whose cuticle is

more irregular. The globular spermatheca connects directly to the oviduct. The

oviduct widens to form a short vagina into which the accessory female glands

open. The female aperture is slightly posterior to the male aperture.

Discussion

The single specimen is similar to P. meckelii, but the shape of the penial

stylet differs (Fig. 1C, 2B), and the vagina is longer in P. meckelii (Fig. 2C).

Pleurobranchaea bonnieae sp. nov.

Figs 1D, 13-14

Type material

Holotype: United States National Museum 809999, R. V. Gerda, Station G

1001, Florida (27°N, 80°W), 61 m, 21 May 1968, one dissected specimen and

slides of the jaws, radula and genital organs.

Etymology

This species is named for Bonnie J. Gosliner.

30 ANNALS OF THE SOUTH AFRICAN MUSEUM

ol Ny

Fig. 13. Pleurobranchaea bonnieae sp. nov. A. Dorsal view of preserved animal, scale

5mm. B. Abnormal rows of radula. C—D. Jaw platelets, scale 0,03 mm.

Description

The preserved animal is 15,5 mm long and 7 mm wide. Its mantle is slightly

tuberculate, white, with bits of pigment retained on the rhinophores and the

mantle. The foot is shorter than the notum anteriorly and longer posteriorly

(Fig. 13A). The contracted oral veil is short and smooth. On the hind end of the

foot is a prominent spur. A pedal gland was not observed. The gill rachis is

smooth with about twenty pinnules on either side. The prebranchial gland opens

at the beginning of the rachis. The genital apertures are retracted and there ap-

pears to be a single small pore without a flap.

The colourless jaw plates are 3 mm long and 1,5 mm wide. The platelets

are up to 0,06 mm high. They are arranged in transverse, slightly slanting rows.

Their surface is hexagonal. Near the anterior end they are about 0,034 by

0,018 mm (Fig. 13C), while posteriorly they are 0,046 by 0,018 mm (Fig. 13D).

FAMILY PLEUROBRANCHAEIDAE Sil

A ee. ite

2 e oe a0)

6 PRI es ~

A <<: $e ene SE . oy

Le. . .

RAO

state ses cectetetet one Sess

eee el lalate ee

—. 35

Fé a2 2 =

: . 5 0 off .

% . o . . . ‘4 .

° rae TH °

Sr EO Ae

D eo°

ie 0° e

g oe

7 fe

PS SSE

BOSS

: afl 7 Soot?

: Oe OS -.

Ceo ee eae OR Sica

DOA ite tie eerturires rere

=e D

Fig. 14. | Pleurobranchaea bonnieae sp. nov. A. Penial sac. B. Female organs. C. Optical

transverse section of penis. D. Root of penis. E-G. Parts of penis in sheath, scale 0,2 mm.

32 ANNALS OF THE SOUTH AFRICAN MUSEUM

The platelets bear nine to twelve denticles. The radula is 4,5 mm long and

2,4 mm wide. The largest of the light yellow, bicuspid teeth is 0,36 mm long.

The formula is 35 X 55.0.55. About six of the outermost teeth are unicuspid. In

the stomach was a 2 mm long aeolid nudibranch.

The reproductive organs were torn during preparation, but the male organ

is complete. The globular penial sac is 1,2 mm in diameter, the atrium 1,6 mm

long (Fig. 14A). A narrow efferent duct, 0,05 mm wide, traverses the penial sac

and continues into the 0,14 mm wide penis proper. The penis is contained within

a 0.25 mm wide penis sheath (Fig. 14D). For the first 1,4 mm of the looping

penis the lumen is closely sinuous (Fig. 14E). Beyond this point the diameter of

the penis diminishes to 0,07 mm while the sheath narrows to 0,12 mm in di-

ameter (Fig. 14F). The 20 mm long organ is so tightly coiled that its further

structure is not recognizable, except for the transverse section (Fig. 14C) and

the tip of the penis (Fig. 14G). As a 0,082 mm wide tube the penis opens

through a tiny papilla into the fundus of the atrium. On the basis of the structure

of the penis and wide sheath around it we guess that the sheath is everted for

copulation and the delicate, non-cuticular penis is protruded.

Discussion

While the single specimen may be juvenile, and the adult may have a cuti-

cular penis, P. bonnieae can still be separated from all described members of the

genus. P. bonnieae is most similar to P. vayssierei (Fig. 19B) and P. notmec

(Fig. 18F), but differs by the wide base of the penis and in its shape in transverse

section (Fig. 14C).

Pleurobranchaea bubala sp. nov.

Figs 15-17

Type material

Holotype: SAM-—A35231, 1 specimen and egg mass, Buffels Bay at Cape

Point, 10 m, collected by William R. Liltved, 4 October 1981.

Paratypes: SAM—A33966, one specimen, Simonstown harbour, Simon’s

Bay; SAM-—A35232, 2 specimens, The Mill, Bakoven, Cape Peninsula, 20 m,

collected by T. M. Gosliner, 16 September 1982.

Other material

SAM-—A35233, 4 specimens, Inhaca, Mozambique, collected by W. Mac-

nae.

Distribution

Specimens are known from the Atlantic coast of the Cape Peninsula to In-

haca, Mozambique.

FAMILY PLEUROBRANCHAEIDAE 33

Fig. 15. | Pleurobranchaea bubala sp. nov., living animal.

Etymology

This species is named bubala after the genus Bubalus, the buffalo, as it is

common at Buffels Bay, Cape Point.

Description

One living animal (Fig. 15) was 70 mm in length. The notum has a pattern

of dark brown pigment and irregular whitish blotches. A well-preserved animal

measured 50 x 35 mm. Its notum extends over the foot on both sides (Fig.

16A-B). Posteriorly it gradually merges with the foot. The oral veil has a row of

small tubercles. A spur is absent. A triangle of pedal glands was visible in one

living specimen. The gill has a smooth rachis with twenty-six pinnae alternating

on either side.

The hexagonal jaw platelets have four to fourteen denticles, rarely more

than ten. The radula formula is 35 x 100.0.100; 20-30 of the outer laterals do

not have a secondary cusp (Fig. 16D).

The genital apertures are protruded and have a slight flap on the hind bor-

der. The glandular tubes of the prostate are united, forming globular masses

(Fig. 16G). The efferent duct between the prostate and penial sac is thick,

0,6 mm in diameter, its lumen is 0,2 mm in diameter. The penial sac (s)

34 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 16. Pleurobranchaea bubala sp. nov. A. Dorsal view of preserved animal. B. Ventral

view of the same. C. Radular tooth from middle of half-row. D. Outermost teeth of row.

E. Spawn. F. Eggs in spawn. G. Reproductive organs.

FAMILY PLEUROBRANCHAEIDAE 35)

Fig. 17. | Pleurobranchaea bubala sp. nov., egg mass.

accompanies the efferent duct (e) inward along the retractor muscle. It is wider

in the middle than in P. tarda, but less globular than that of P. inconspicua. The

cuticular stylet (c) is similar to that of P. tarda and about 5 mm in length. The

atrium (a) is strongly muscular, 6-9 mm long.

The epithelium of the oviduct forms pouches, the spermatocyst (y). From

the spermatocyst the oviduct narrows until it branches to the unstalked bilobed

spermatheca (Fig. 16G). From this point the vagina (v) or insemination duct is

much wider and has a high, folded epithelium. Near the female aperture it re-

ceives the accessory glands and forms a small caecum.

The egg mass (Figs 16E, 17) is a coiled tube of several irregular whorls.

There are one to three eggs per capsule (Fig. 16F).

This species has been observed to feed on other opisthobranchs including

members of its own species.

Discussion

The reproductive organs differ from the other species by the shape of the

penial sac, the double spermatheca and the long glandular vagina. P. bubala is

sympatric with P. tarda in South Africa and is distinguished by its larger size,

abundant white blotches, elongate vagina, and the shape of the prostate (Figs

5A, 16G).

36 ANNALS OF THE SOUTH AFRICAN MUSEUM

Pleurobranchaea notmec sp. nov.

Figs 1E-G, 18

Type material

Holotype: University of Israel, Department of Zoology 1005, eastern Medi-

terranean, off Turkey, 11-140 m.

Paratypes: University of Israel, Department of Zoology, two specimens,

Haifa Bay; one specimen, Palmahim, 80 m; one specimen, No. 16399, northern

Sinai, 45 m.

Etymology

The name refers to the fact that this species was originally identified as P.

meckelii, from which it is distinct.

Description

The size of the preserved animals is from 12 to 40 mm long and 8 to 15 mm

wide.

The surface of the mantle is slightly bossed, without tubercles.

The colour is not preserved, except some pigments in the furrows between

the bosses. The animals are ‘museum’ brown. The notum is smaller than the

foot. Its right border extends out over the gill. The anterior border of the notum

forms the oral veil bearing a row of papillae. The sides of the veil are produced

into pointed cephalic tentacles with split sides. The rhinophores are inserted be-

tween the tentacles and the lateral mantle border. They are as long as the tenta-

cles, blunt, cylindrical, and rolled.

The anterior end of the foot is divided by a transverse furrow. The hind end

bears a pointed dorsal spur. On the ventral side the pedal gland forms a longi-

tudinal furrow.

The gill is about one-third of the body length and has twenty to thirty alter-

nating pinnules on each side. Its anterior three-fourths to four-fifths are adnate.

The anus is dorsal to the middle of the gill. The rachis is smooth.

The prebranchial glands open slightly anteriorly to the gill. A little more

anteriorly are the genital apertures, united by a circular fold. Their shape varies

according to contraction. A flap is absent. The large pharynx often protrudes

from the mouth opening. The labial cuticle bears scattered papillae. The ante-

rior border of the jaws appears whitish on the everted proboscis but is light

brown when cleaned with potassium hydroxide. They are composed of columnar

rodlets of variable shape, reaching 0,2 mm in height. The rodlets are polygonal,

longer than wide. Ectally they measure about 0,05 mm in length and 0,04 mm in

width. Posteriorly they are longer and narrower, the longest attaining 0,11 mm

in length and 0,017 mm in width. Their frontal borders overlap and bear many

small denticles, which are also present on the surface behind the border. The

bases of the rodlets are more uniform in size. Anteriorly they are 0,05 by 0,03

mm; posteriorly 0,07 by 0,03 mm.

FAMILY PLEUROBRANCHAEIDAE 37

nial eZ

MAA SD,

(

ae \

TT WSN

tea WI NL

i ima)

é ian

aT |

Cas

=5 \

——— ss

—————SS——

—

S—-s

———S

SSS

Fig. 18. Pleurobranchaea notmec sp. nov. A. Radular tooth from middle of half-row.

B. Outer radular tooth. C-—E. Optical sections of penis. F. Male reproductive organ, scale

0,3 mm.

HY |

: i

The radula of the largest specimen is 16 mm long; each half is 5 mm broad.

The formula is 50 x 80.0.80. The teeth have a small secondary cusp (Fig. 18A).

In the middle of the half-row the longest teeth attain 0,65 mm in length and dim-

inish gradually towards the outer edge. One to four outer teeth lack a secondary

cusp (Fig. 18B).

38 ANNALS OF THE SOUTH AFRICAN MUSEUM

The genital ducts are so compact that they could not be disentangled. The ovo-

testis follicles contain sperm or oocytes. The slender hermaphroditic duct widens

into a long ampulla, narrows again, and divides into the male and female ducts.

The male duct enters the round prostate (Fig. 18F, q) and emerges as a very thin

efferent duct (e) which penetrates the penial sac (s) near its outer end. Inside

the sac the penis is a long, winding tube, 0,17 mm in diameter. Its lumen is

0,10-0,14 mm in diameter. The penis has a very thin cuticle and is surrounded

by a thin penial sheath (Fig. 18C—E), which must evidently be everted. After

forming about six loops, the penis straightens and emerges in the male atrium as

a small papilla (Fig. 18F). The atrium (a) is a wide, thick-walled sac that opens

in front of the female pore.

It was not possible to follow the oviduct. It did not seem to have a spiralled

spermatocyst.

Discussion

The present animals from Israel were originally identified as Pleurobran-

chaea meckelii. However, P. meckelii differs by its long whip-like, cuticular

penial stylet. The single specimen of P. vayssierei (Fig. 19B) is also similar but

lacks a cuticle around the efferent duct in the penial sheath.

Pleurobranchaea vayssierei sp. nov.

Fig. 19

Type material

Holotype: Museum National d’Histoire Naturelle, Paris, from Algiers, 1900,

one specimen among Vayssiére’s specimens of P. meckelii, No. 52. (See p. 11.)

Etymology

The name refers to the fact that the unique specimen was found in the col-

lection of Albert Vayssiere.

Description

The preserved animal is 32 mm long, 11 mm broad and 10 mm high. The

notum is as large as the foot, smooth and colourless. There is a tiny spur on the

tip of the foot (Fig. 19A), and a weak metapodial gland on its underside. The

oral veil is smooth, perhaps as a result of 82 years of preservation. The gill has a

smooth rachis with twenty-six pinnae. The jaw plates (Fig. 19C) are broken.

Their platelets are relatively broad (Fig. 19D). The platelets bear up to fifteen

denticles, which disappeared after mounting them in glycerine. The radular for-

mula is 35 x 65.0.65. The teeth are dark greyish and bicuspid (Fig. 19E). None

are unicuspid on the outer border.

The male duct forms many loops in the tight transparent penial sac (Fig.

19B). A stylet is not recognizable. The single specimen was so fragile that the fe-

male organs could not be dissected completely. The spermatocyst is a slight

FAMILY PLEUROBRANCHAEIDAE 39

ef / Ci —. 5)

Cali (i we

A\t

D F

Fig. 19. Pleurobranchaea vayssierei sp. nov. A. Preserved specimen from Algiers. B. Penial

sac. C. Anterior view of pharynx with jaws. D. Jaw platelets. E. Radular teeth from middle

of half-row. F. Twenty-second outer tooth. G. Five outermost teeth.

widening of the oviduct, and the oviduct is continued into a long, rather narrow,

ciliated vagina.

Discussion

The most similar species is P. bonnieae from Florida (Fig. 14A), which has

an enlarged root of the penis. In P. notmec up to twenty teeth of the radula are

unicuspid.

40 ANNALS OF THE SOUTH AFRICAN MUSEUM

Pleurobranchaea (Macfarlandaea) subgen. nov.

Type species

Pleurobranchaea californica MacFarland, 1966.

Diagnosis

Pleurobranchaeidae with rudimentary secondary cusps on all radular teeth.

Pleurembolic penis with cuticular stylet.

Etymology

The subgeneric name honours the late Frank Mace MacFarland of Stanford

University.

Pleurobranchaea (Macfarlandaea) californica MacFarland, 1966

Figs 20-22

Pleurobranchaea californica MacFarland, 1966: 94-101, pl. 15 (figs 16-28), pl. 17 (figs 1-17).

Chivers, 1967: 515-521.

Pleurobranchaea sp. Coan, 1964: 173.

Fig. 20. Pleurobranchaea (Macfarlandaea) californica MacFarland, 1966, living animal.

FAMILY PLEUROBRANCHAEIDAE 41

Fig. 21. Pleurobranchaea (Macfarlandaea) californica MacFarland, 1966 (after MacFarland

1966). A. Reproductive organs (retracted). B. Radular tooth. C. Reproductive organs with

everted penis.

Distribution

California, from Klamath River to San Diego, 10-470 m.

Description

The living animal (Fig. 20) is brown with white blotches, up to 335 mm

long. The rhinophores are typical of the family Pleurobranchaeidae. The jaw

platelets are columnar with denticles. The radular formula of a 165 mm long ani-

mal is 52 x 130—145.1.130-145. All the lateral teeth have a rudimentary second-

ary cusp (MacFarland 1966: pl. 15 (figs 22—24)). The penis sac (= penis sheath,

MacFarland 1966) is long (2 cm in an adult specimen) and not quite close

around the penial sheath (= preputium, MacFarland 1966). The efferent duct

accompanies the retractor muscle upwards and they enter the sheath together.

Here the sheath separates from the whip-like muscular penis. The egg mass is a

highly convoluted collar (Fig. 22).

Coan (1964: 173) observed that the species is carnivorous and cannibalistic.

42 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 22. Pleurobranchaea (Macfarlandaea) californica MacFarland, 1966. Egg mass.

Euselenops Pilsbry, 1896

Pleurobranchaea (Euselenops) Pilsbry, 1896: 228. Thiele 1931: 419.

Neda Adams & Adams, 1854. Non Mulsant.

Oscaniopsis Bergh, 1897: 53. Vayssiére, 1901: 6.

Type species

Pleurobranchus luniceps Cuvier, 1817.

Diagnosis

Notum much smaller than foot, penis papillate.

Euselenops luniceps (Cuvier, 1817)

Fig. 25D-G

Pleurobranchus luniceps Cuvier, 1817: 186.

Oscaniopsis semperi Bergh, 1897: 55, pl. 6 (figs 7-27).

Oscaniopsis compta Bergh, 1897: 58, pl. 8 (figs 11-27).

Oscaniopsis amboinei Vayssiére, 1898: 9, fig. 27; 1901: 15, figs 190-204.

Non Oscaniopsis pleurobranchaeana Bergh, 1907: 35, pl. 4 (figs 16-21).

Oscaniopsis luniceps Vayssiére, 1901: 15.

Euselenops luniceps O’Donoghue, 1929: 55, figs 65-74. Pruvot-Fol, 1933: 107. Burn, 1962: 131.

Lin Guangyu & Tchang-Si, 1965: 269, 275, fig. 4.

FAMILY PLEUROBRANCHAEIDAE 43

Distribution

Indo-Pacific: Hawaii, China, Hong Kong, Hainan, Japan, Philippines, Am-

boina, Queensland, Mauritius, Inhaca, South Africa, ?Red Sea.

Description

The length is up to 90 mm (Lin Guangyu & Tchang Si 1965) with the notum

much smaller than the foot and its hind end forming a projecting tip. The radu-

lar teeth are unicuspid. The prostatic cells are contained in the efferent duct and

the acrembolic penis is covered with soft cones.

Discussion

One 12 mm long specimen from Palmahim, Israel, had the outer aspect of

Euselenops with a broad velum, small notum, large foot, and a notch in the hind

end of the mantle. At first glance it seemed to be a Euselenops sp., but it was

discovered that the radula was bicuspid, that the notal notch was the siphon of

the gill, and that there was a large spur on the tip of the foot (Fig. 25B), indicat-

ing that it was a Pleurobranchaea sp. As the reproductive organs were not devel-

oped, the species could not be determined.

Pleurobranchella Thiele, 1925

Pleurobranchella Thiele, 1925: 283. Willan, 1977: 151.

Pleurobranchoides O’Donoghue, 1929: 62.

Pleurobranchaea (Euselenops) Pleurobranchella Thiele, 1931: 419.

Type species

Pleurobranchella nicobarica Thiele, 1925.

Diagnosis

Unicuspid species of Pleurobranchaeidae; notum covering foot on all sides;

acrembolic penis with cuticular hooks.

Pleurobranchella nicobarica Thiele, 1925

Figs 23-24

Pleurobranchella nicobarica Thiele, 1925: 283, pl. 33 (figs 9-11).

Pleurobranchoides gilchristi O’Donoghue, 1929: 62, figs 75-85. Eales, 1937: 371.

Pleurobranchoides sp. Eales, 1938: 90, fig. 10.

Material

SAM-A33973, three specimens (105 mm, 115 mm, 150 mm long, pre-

served), Monte Belo, Mozambique, 400 m.

Distribution

Nicobares, Gulf of Aden, southern Africa, 220-269 m.

44 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 23. | Pleurobranchella nicobarica Thiele, 1925. A. Diagram of reproductive organs.

B. Dorsal aspect. C. Radular tooth. D. Penial hook, scale 100 um. E. Ventral aspect.

Description

The large mantle extends far over the foot on all sides (Fig. 23E). The radu-

lar formula is 60 x 133.0.133. The extruded penis (Fig. 24B) consists of a wide

everted atrium and a long cylindrical, slightly spiralled penis proper. Its concave

FAMILY PLEUROBRANCHAEIDAE 45

Fig. 24. Pleurobranchella nicobarica Thiele, 1925. A. Outer end of penis, scale 2,0 mm.

B. Extruded penis, scale 5,0 mm. C. Penial hooks. D. Penis removed from retracted sac.

side is smooth; the convex side is covered with hook-shaped papillae (Fig.

24C-D). The extruded atrium is 30 mm long and 8 mm wide in the largest speci-

men. The penis proper is 7 by 2 mm in the largest specimen and 6 by 2,5 mm in

the smallest.

Discussion

Though Thiele (1925) did not figure the penis of his Pleurobranchella nico-

barica, his description is precise enough to identify the present specimens with

his material. O’Donoghue gave only a short description of Pleurobranchoides

46 ANNALS OF THE SOUTH AFRICAN MUSEUM

gilchristi, but it is considered sufficient to synonymize it with Pleurobranchella

nicobarica.

Eales’s (1938: 90, fig. 10) Pleurobranchoides sp. from the Gulf of Aden has

a papillate penis and is certainly identical with Pleurobranchella nicobarica.

Koonsia Verrill, 1882

Koonsia Verrill, 1882: 545.

Type species

Koonsia obesa Verrill, 1882.

Remarks

Pilsbry (1896: 221) considered Pleurobranchillus Bergh, 1892, as a synonym

of Koonsia, but Bergh (1897) transferred Pleurobranchillus to Pleurobranchaea,

as did Willan (1977: 153).

Koonsia obesa Verrill, 1882

Fig. 25C

Koonsia obesa Verrill, 1882: 545, partim; 1884: pl. 28 (fig. 7); 1885: 571, fig. 107. Abbott, 1974:

349, fig. 2046.

Distribution

Western Atlantic, 7Martha’s Vineyard: ?Delaware Bay, 400-560 m.

Discussion

Verrill (1882) included two distinct species in his original description. In his

description of the genus Verrill (1882: 545) stated that ‘the back is overhanging

both on the sides and posteriorly, and there is a distinct mantle edge all around’.

This agrees with the species description that ‘the mantle forms a ring along the

lateral and posterior borders’; it does not correspond to the figure of Koonsia

obesa but to his distinct paratypes, which have been examined (see P. confusa).

He stated that ‘the foot is narrower’, as in the paratype. In the generic diagnosis

Verrill stated that ‘the verge is armed with small hooks, but the spicule, present

in Pleurobranchaea, is not protruded in any of our specimens of Koonsia, if

present’. He neglected to mention this in the species description, but noted of

Pleurobranchaea tarda ‘the verge . . . with rows of minute recurved hooks near

the end, and terminated by a slender, curved spicule’ (Verrill 1882: 546).

The paratypes without hooks seen by Bergh (1897), Vayssiére (1901), and

the authors (Fig. 3) are quite different from Verrill’s figure (1885: 107). They

also differ from his figures of P. tarda (1882, pl. 58 (fig. 6); 1885, fig. 105), which

possess a stylet, but lack hooks.

As Bergh (1897: 33) did not find penial hooks in the paratype specimen of

‘Koonsia obesa’ from the original locality, he rejected Koonsia and placed K.

obesa in Pleurobranchaea. The authors believe that the distinct penial spines

FAMILY PLEUROBRANCHAEIDAE

? £

(<)

fe)

fo)

os]

flo:

Soo

poe ove6

fe 250800909

5 OO a 00000

Grn 00 “50 RCocs

Fig. 25. A. Gigantonotum album Lin Guangyu & Tchang Si, 1965, preserved specimen (after

Lin Guangyu & Tchang Si 1965). B. Juvenile Pleurobranchaea sp. from Israel. C. Koonsia

obesa Verrill, 1882, holotype (after Verrill 1882). D. Euselenops luniceps (Cuvier, 1817) re-

productive system (after Vayssiére 1901). E. Pleurobranchaea maculata (Quoy & Gaimard,

1832), preserved specimen (after Baba 1937). F—G. Euselenops luniceps. F. Penial papillae

G. Whole animal (after Lin Guangyu & Tchang Si 1965).

(after Bergh 1897).

48 ANNALS OF THE SOUTH AFRICAN MUSEUM

described by Verrill indicate that his type material actually represents two

species and therefore name the paratype examined Pleurobranchaea confusa. If

the slender species (Fig. 25C) is found again, its generic placement depends

upon whether the radula is unicuspid or bicuspid. Until such time it is preferred

to maintain Koonsia as a distinct genus. Pleurobranchella nicobarica Thiele,

1925, Euselenops luniceps (Cuvier, 1817), and Pleurobranchus hirasei Baba,

1971, are the only other notaspideans known to possess an ornamented penis.

Gigantonotum Lin Guangyu & Tchang Si, 1965

Gigantonotum Lin Guangyu & Tchang Si, 1965: 270.

Type species

Gigantonotum album Lin Guangyu & Tchang Si, 1965.

Diagnosis

Pleurobranchaeid with a very wide mantle that covers the foot and the gill

completely. Its borders are almost half as wide as the foot. Its underside bears

nodules. The radula is unicuspid; its formula is 52 x 144.1.144.

Gigantonotum album Lin Guangyu & Tchang Si, 1965

Fig. 25A

Gigantonotum album Lin Guangyu and Tchang Si, 1965: 270, figs 5—6, pl. 1 (fig. 5d, v).

Pleurobranchella alba Willan, 1977: 153.

Distribution

South of Hainan Island, China Sea, 220 m.

Discussion

Lin Guangyu & Tchang Si (1965) compared their species with Koonsia, but

stated that in Gigantonotum the foot is covered by the mantle, while it extends

posteriorly for a considerable distance in Koonsia.

On the whole, Gigantonotum is similar to Pleurobranchella which, how-

ever, lacks a rachidian tooth. The geographic proximity makes a synonymy

probable. Willan (1977: 151) synonymized Gigantonotum with Pleurobranchella,

but considered only the incomplete previous descriptions. As long as the repro-

ductive organs of G. album remain unknown it is preferable to consider Gigan-

tonotum as a distinct but doubtful genus.

ZOOGEOGRAPHY OF THE PLEUROBRANCHAEIDAE

Only preliminary zoogeographical remarks can be made, owing to the fact

that many records are based on unrecognizable, incompletely described, or un-

justly synonymized species. In other cases information about the fauna of a re-

gion is incomplete. For example, Edmunds (1977) did not record any nota-

spideans from Ghana.

FAMILY PLEUROBRANCHAEIDAE 49

Five species of Pleurobranchaeidae have been recorded from the western

Atlantic: Pleurobranchaea tarda, P. agassizii, P. inconspicua, P. bonnieae, and

Koonsia obesa. Of these, P. tarda and P. inconspicua are more widespread. P.

tarda is known from the eastern Atlantic coast of Africa from Ghana and from

South Africa, where it extends into the Indian Ocean. P. inconspicua is also

known from the Mediterranean and west Africa. Pleurobranchaea meckelii, P.

notmec and P. vayssierei are known only from the Mediterranean, while P. au-

gusta has been recorded only from west Africa. P. bubala is known from the At-

lantic and Indian Ocean coasts of southern Africa.

Pleurobranchaea brockii and Pleurobranchella nicobarica are both known

from the eastern and western Indian Ocean. Euselenops luniceps is widespread

in the Indo-Pacific where it has been recorded from Hawaii to South Africa.

Other species known from the Indo-Pacific region—Pleurobranchaea maculata

and Gigantonotum album—may be widespread but their ranges need to be es-

tablished from reliable records.

Pleurobranchaea californica is restricted to the Pacific coast of North

America.

ACKNOWLEDGEMENTS

We are deeply indebted to our colleagues at other institutions who gener-

ously provided assistance and specimens. To Frederick M. Bayer, Harald Reh-

der, and Joseph Rosewater of the United States National Museum, Washington;

Alexander Barash, University of Israel, Tel Aviv; Hanne Just, Zoology Mu-

seum, Copenhagen; Philippe Bouchet, Museum National d’Histoire Naturelle,

Paris; and Luise Schmekel, Zoologisches Institut der Universitat, Minster, we

extend our sincere thanks.

Several staff members of the South African Museum provided considerable

assistance. William Liltved prepared the final figures and collected specimens.

Michelle van der Merwe printed the photographs and Marcelle Scheiner typed

the final drafts of the manuscript. To them we also extend our appreciation.

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Marcus, Er. & Marcus, Ev. 1968. Some opisthobranchs from Ivory Coast. Bull. Inst. fr. Afr.

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FAMILY PLEUROBRANCHAEIDAE 51

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ABBREVIATIONS

a atrium n female pore

b_ base of penis O spermoviduct

Cc penial stylet p penis

d hermaphrodite duct q_ prostate

e efferent duct r retractor muscle

f oviduct Ss penial sac

g mucous gland t spermatheca

h_ penial sheath Vv vagina

i root of penis w body wall

j rhinophore xX spur

k gill y spermatocyst

1! albumen gland z foot gland

m ampulla ? female gonopore

6 male gonopore

y ye A

ae ee

- y -_ ares

‘

)

}

6. SYSTEMATIC papers must conform to the International 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 namé 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

>] 6

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. Du Toit 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.

EVELINE d. B.-R. MARCUS

TERRENCE M. GOSLINER

REVIEW OF THE FAMILY

PLEUROBRANCHAEIDAE

(MOLLUSCA, OPISTHOBRANCHIA)

ANNALS

OF THE SOUTH AFRICAN

MUSEUM

CAPE TOWN :

INSTRUCTIONS TO AUTHORS

. MATERIAL should be original and not published elsewhere, in whole or in part.