Tropical Zoologs 2 : 145- 157, 1989

Kalyptorhynchia (Plathelminthes ) from the Kenyan Coast, with descriptions of four new species

P.E.H. J o u k an d A.J.-P. D e V o c h t Research Group Zoology, Department SBM, Limburgs Universitair Centrum, B-3610 Diepenbeek, Belgium

Received 6 March 1989, accepted 1 ]une 1989

Sandy sediments and algae from the litoral and sublitoral zone of the coast near Mombasa were sampled. Eighteen species of were collected, of which four new species of Schizorhynchia are described here, two Cheliplana species, one Baltoplana sp. and one Carcharodorhynchus sp. Cheliplana asica terminalis Brunet 1968 described from the Mediterranean and also recorded from Somalia was found as well. Five out of 13 species of Eukalyptorhynchia were already recorded from Somalia. Polycystis ali Schockaert 1982, P. subcontorta Schockaert 1982 and Cincturorhynchus karlingi Schockaert 1982 were newly described from the Somalian coast. P. contorta Schockaert & Karling 1975 also known from Norway and the Mediterranean and Gyratrix hermaphroditus Ehrenberg 1831 is an ubiquist. Six new species of Eukalypto­ rhynchia collected from algae will be described in a later publication. Additional information is given on Alcha evelinae Marcus 1949 and Toia ycia Marcus 1952.

k e y w o r d s : Plathelminthes, Rhabdocoela, Kalyptorhynchia, Kenya, new species, species list.

Introduction ...... 146 Material and methods ...... 146 Descriptions ...... 146 Eukalyptorhynchia...... 146 Polycystis ali Schockaert 1982 146 Polycystis contorta Schockaert & Karling 1975 147 Polycystis subcontorta Schockaert 1982 147 Alcha evelinae Marcus 1949 147 Gyratrix hermaphroditus Ehrenberg 1831 148 Cincturorhynchus karlingi Schockaert 1982 148 Toia ycia Marcus 1952 148 Schizorhynchia...... 149 Carcharodorhynchus involutus n. sp...... 149 Baltoplana bisphaera n. sp. 149 Cheliplana asica terminalis Brunet 1968 151 Cheliplana pileola n. sp...... 151 Cheliplana textilis n. sp...... 153 146 P.E.H. Jouk and A.J.-P. De Vocht

Abbreviations ...... 155 Acknowledgements ...... 155 References ...... 156

INTRODUCTION

Up to now, records on marine Turbellaria from Africa are very few and solely restricted to the Somalian coast(Schockaert 1971, 1982; Schockaert & M artens 1985, 1987). Thusfar,Schockaert (1971, 1982) recorded 10 species of Kalyptorhyn­ chia, seven of which were new to science. In this first contribution of Kenyan Turbellaria, 12 species of Kalyptorhynchia are listed, of which four new species of Schizorhynchia are described here. Six new species of Eukalyptorhynchia will be described in a later publication.

MATERIAL AND METHODS

All the species were collected by the first author from marine habitats in the neighbourhood of Mombasa in September-October 1985 and May-June 1987. Living were extracted from the sediment with a MgCk-solution isotonic to seawater (M a r t e n s 1984). Species living on algae were separated by stirring the algae vigorously in seawater. After studying the animals alive, whole mounts were made with lactophenol or polyvinyllactophenol. If found in sufficient numbers, some specimens were fixed in Bouin’s fluid and serially sectioned. Sections were stained with Heiden- hain’s iron hematoxylin, using eosin as counterstain. All material, included types, is deposited in the collection of the Limburgs Universitair Centrum ( = LUC).

DESCRIPTIONS

Eukalyptorhynchia

Polycystis ali Schockaert 1982 (Fig. 3A) Distribution. Somalia, N of Mogadiscio (Hawadli) on algae in pools on the rocky shore at low tide (Schockaert 1982). California (USA) in tidal pools and shallow water in gravel, on stones and algae(K a r lin g 1986). Galapagos Islands, in a tidal pool (see K a r lin g 1986). Locality in Kenya. At the mouth of Tudor Creek, Mombasa (McKenzie Point), onThalassia hemprichii, covered by the epiphyteEnteromorpha kylinii in pools on the rocky shore at low tide.

Material. One studied alive and mounted.

Additional information. K a r lin g (1986) distinguishes three «forms» based on the differences in shape and size of the stylet. These «forms» match the populations from the three areas of distribution. The stylet in our specimen is 20 pm high and 41 pm wide distally, which is somewhat larger than inSchockaert’ s material (17 pm high Kalyptorhynchia from Kenya 147

and 28 pm wide). The collar of the stylet has a finely toothed edge all around and shows no slit (Fig. 3A). These characters indicate that this specimen belongs to the «Somali» form.

Polycystis contorta Schockaert & Karling 1975 Distribution. Norway, S of Bergen, in shell gravel, 7-50 m deep. Mediterranean, Marseille area (France) inAmphioxus- sand (Schockaert & Karling 1975). Somalia, N of Mogadiscio (Hawadli) on algae in pools on the rocky shore at low tide (Schockaert 1982). Locality in Kenya. Same locality as forP. ali. Material. One animal studied alive and mounted. Additional information. The length of the stylet in our specimen (45 pm, mea­ sured along its central axis) is only slightly less than the lengths (51-53 pm) observed by Schockaert & Karling (1975). The drawing ofSchockaert (1982) shows a smaller stylet (about 40 pm) for his Somalian specimens. These observations indicate the presence of a smaller stylet in the East African populations.

Polycystis subcontorta Schockaert 1982 Distribution. Somalia, N of Mogadiscio (Hawadli) on the sandy bottom of a pool on the rocky shore at low tide. Locality in Kenya. Same locality as forP. ali.

Material. One animal studied alive and mounted. Additional information. The stylet in the Kenyan specimen is 76 pm long (measured along the central axis of the stylet). This is much smaller than in the Somalian specimens (126 pm long).Schockaert (1982) mentions the presence of two optically empty vesicles communicating with the oviducts, but their function re­ mained unknown because of the incomplete female maturity of his specimens. In our specimen, these vesicles contained sperm. This is the first recording of seminal receptacles in the genusPolycystis.

Alcha evelinae Marcus 1949 (Fig. 3B) Distribution. Brazil, on seaweed in the eulitoral zone(M a r c u s 1949). California (USA) in tidal pools in gravel and on seaweed(Karling & Schockaert 1977). Locality in Kenya. Same locality as forP. ali and at 6 m depth onThalassia hemprichii, partly covered by the epiphyteSyringodium isoetifolium (English Point).

Material. This species was the most abundant representative of the Eukalyptorhynchia in the area investigated. More than 50 specimens were studied alive and at least 25 were preserved as whole mounts.

Additional information. According to the descriptions M of a r c u s (1949) and Karling & Schockaert (1977) our specimens clearly belong to this species. The length of the male cuticular apparatus (Fig. 3B) varies from 33 to 50 pm. In the Brazilian material, it was 40 pm long, in the California 53 pm. 148 P.E.H. Jouk and A.J.-P. De Vocht

Gyratrix hermaphroditus Ehrenberg 1831 Distribution. This species has always been considered to be a cosmopolitan and holeuryhaline species. For example, it has been encountered on the European Atlantic and North Sea coasts(B o a d en 1963,K a r lin g 1963, Ax & Ax 1970,S c h ilk e 1970), in fresh water in Finland and Germany(H e itk a m p 1978), in the Baltic (K a r lin g 1963, 1974), the Mediterranean( B r u n e t 1980, Puccinelli & Curini-Galletti 1987), the Black Sea (Magic Fiha 1968), Hawaii (K a r lin g et al. 1972), California (Karling & Schockaert 1977), Bermuda(K a r lin g 1978), NE Asia (E v d o n in 1971) and Somalia (Schockaert 1982). Locality in Kenya. At the mouth of Tudor Creek, Mombasa (English Point), at 6 m depth onThalassia hemprichii, partly covered by the epiphyteSyringodium isoetifo­ lium. Material. One animal studied alive and mounted. Additional information. The measures of the cuticular apparatus are: stylet, 90 pm; sheath, 28 pm; stalk, 44 pm. The range of variation in size of the cuticular elements inG. hermaphroditus is enormous; respectively 75-275 pm, 18-87 pm and 42-160 pm (Karling & Schockaert 1977, K a r lin g 1978). Recently, karyological and morphological research have reported thatG. hermaphroditus is a species complex (H e itk a m p 1978, L ’H a r d y 1986, Puccinelli & Curini-Galletti 1987). Thus the classification of our specimenG. as hermaphroditus is preliminary.

Cincturorhynchus karlingi Schockaert 1982

Distribution. Somalia, N of Mogadiscio (Hawadli) on the sandy bottom of a pool and on algae on the rocky shore at low tide. Locality in Kenya. Same locality as forP. ali.

Material. One animal studied alive (and fixed).

Additional information. In conformity with the observationsS c of h o c k a e r t (1982), our specimen is coloured with a reddish brown pigment and possesses a proboscis about 1/5 of the total length. The topography of the genital organs in Schockaert (1982) is valid for this specimen as well, except for the terminal bursa which could not be observed.

Toia ycia Marcus 1952

Distribution. Brazil, on seaweed in the upper-eulitoral(M a r c u s 1952). Nether­ lands, Grevelingen onZostera marina, 2.5 m deep (Martens & Schockaert 1981). Locality in Kenya. Same locality as for P.ali.

Material. One animal studied alive and mounted. Additional information. Length of cirrus: 32 pm (needles 1-2 pm). The specimen found byMartens & Schockaert (1981) had a much larger cirrus (67 pm long, needles 3.8 to 4.8 pm).M a r c u s gives no measures for the cirrus, but from his drawings, a length of approximally 45 pm can be deduced. Kalyptorhynchia from Kenya 149

Schizorhynchia

Carcharodorhynchus involutus n. sp. (Figs 1A-C and 3C-D) Locality in Kenya. At the mouth of Tudor Creek, Mombasa (English Point), in clean, fine sand at mid tide level (type locality). Material. Two animals studied alive, one mounted (holotype LUC No. 140) and one serially sectioned (paratype LUC No. 141). Description. C. involutus is 415-560 pm long (fixed) with a subterminal adhesive girdle but without pigmented eyes. The pharynx rosulatus (diameter 70 pm) is situated in the posterior part of the second third of the body. The proboscis is symmetrical and measures 53-78 pm. Two U-shaped «fields» of small teeth are situated at the inner side of both proboscis lips. Posteriorly they contain four or five rows of teeth, anteriorly only one or two rows. All teeth are uniform (1 to 1.5 pm). The male genital organs consist of a pair of long, folded testes, situated ventro- laterally just in front of the pharynx. The foldings cause the appearance of four or five separate follicles. The two seminal vesicles are located just behind the pharynx. The copulatory bulb (45 pm long) has a very thick muscular wall and contains prostatic secretion. The hard parts of the male copulatory organ consist of a 44 pm long stylet, which encloses a 20 pm long cirrus. The ejeculatory duct forms longitudinal ridges in the stylet. The proximal parts of the median ridges are lamellated, forming the cirrus. The stylet is formed by sclerotisation of the everted part of the ejaculatory duct. It opens by a lateral split in the distal third of the stylet. The stylet lies in a muscular male atrium. The male duct leads ventrally into the common genital atrium, which opens in the genital pore, located in the rearmost eighth of the body. The two ovaries are located dorsally just in front of the common genital pore. The yolk glands form two dorso-median rows in the body, reaching from just behind the brain to the ovaries. In the rear there is a bursa with an accumulation of sperm. Discussion. Based on the general morphology and on the morphology of the proboscis, this species is classified in the genusCarcharodorhynchus. All representa­ tives of this genus are elongated, eyeless and have a symmetrical or asymmetrical proboscis, mostly armed with two fields of teeth at the inner side of the proboscis lips. They have paired gonads (paired ovaries, yolk glands and one to eight pairs of testes). The male copulatory organ is provided with a cirrus or a stylet, sometimes with both structures. C. involutus can easily be distinguished from the other species in this genus by the structure of its proboscis and male copulatory organ. polyorchis Only C. L’Hardy 1963, C. flavidus Brunet 1967, C.ambronensis Schilke 1970 and C. arista Noldt & Hoxhold 1984 have a symmetrical proboscis. All other species have an asymmetrical proboscis.C. involutus is the onlyCarcharodorhynchus species with a large stylet enclosing the smaller cirrus. In the other species of the genus, the stylet is situated within the cirrus.

Baltoplana bisphaera n. sp. (Figs 1D-E and 3E-F) Locality in Kenya. At the mouth of Tudor Creek, Mombasa (English Point), as well in the seepage (see C. textilis) as in the non-seepage area (see C.involutus). pr—

— pr phc

prg

t —

— t V s

mii ^ b— ci s- cb- vs

ov- |:0- md cb

c i -

b - —ov

— p bt-

BCE 20 pm

Fig. 1. — A-C, Carcharodorhynchus involutus n. sp.; A, squeezed live specimen (free-hand drawing); B, proboscis; C, male copulatory organ (B-C from whole mount). D-E, Baltoplana bisphaera n. sp.; D, squeezed live specimen (free-hand drawing); E, proboscis (from whole mount). Kalyptorhynchia from Kenya 151

Material. Two animals studied alive, one mounted and one sectioned (holotype LUC No. 142 and paratype LUC No. 143 respectively). Description. The morphology B. of bisphaera fits the description ofBaltoplana magna Karling 1949 very well. The animal is ahoot 1.7 mm long (measured on the fixed specimen). The pharynx (200 pm long), provided wilh a long cuvily, lies in the anterior third of the body. A caudal haptic girdle is present. The proboscis is 50-51 pm long and 22 pm wide, the hooks have a length of 28-30 pm, the muscle bulbs measure 26-28 pm in length and 9-11 pm in width. The paired testes are situated in front of and beside the cylindrical pharynx, the paired ovaries behind the male copulatory organ. The copulatory organ is approxima­ tely 180 pm long (126-330 pm B. in magna). In the copulatory bulb only the two diverticles (diameter 35-40 pm) are armed with needles. The ductus ejaculatorius is completely unarmed. A muscular duct frontally with two types of glands can be observed laterally, its function is unknown. Discussion. Up to now, the genusBaltoplana contained only two species:B. magna, known from the Baltic(K a r lin g 1949), the Mediterranean (Ax 1956,B r u n e t 1980) and the North Sea (Ax 1956) andB. valkanovi Ax 1959 known from the Black Sea and the Bosporus (Ax 1959). The three species can easily be identified by their different copulatory organs. In B. magna, the longish copulatory bulb contains a cirrus with two distal diverticles, which, like the cirrus, are armed with small cuticular needles(K arling 1949). B. valkanovi has only an armed cirrus, lacking the two diverticles.B. In bisphaera, only the two diverticles are armed. The copulatory organB. of bisphaera is about twice as long as wide (against3 times as long as wide inB. magna). Furthermore,B. valkanovi and B. bisphaera have similar lengths (B. valkanovi reaches maximally 1.8 mm in length), while B. magna has a body length of3 to 4 mm(K arling 1949, Ax 1956). All species ofBaltoplana are encountered in brackish water habitats.

Cheliplana asica terminalis Brunet 1968

Distribution. Mediterranea, Laguna of Brusc, France(B r u n e t 1968) in muddy sand. Somalia, S of Mogadiscio (Djezira) at the inlet of the salt garden in mud (Schockaert 1982). Locality in Kenya. Same locality as for C.involutus. Material. Two animals studied alive, one mounted. Additional information. Cheliplana Marcus asica 1952 was originally described from the Brazilian coast (muddy sand in brackish water). The subspeciesC. asica terminalis differs from C. asica asica by having a slight sclerotisation of the terminal part of the ejaculatory duct and of the wall of the penial papilla protruding in the atrium. These characters were also observed on the Kenyan specimens, and therefore we regard them as C. asica terminalis.

Cheliplana pileola n. sp. (Figs 2A-C and 3G-H) Locality in Kenya. In the mangrove area at the mouth of Tudor Creek, Mombasa (McKenzie Point) in a tidal pool with very fine sand, mixed with some detritus. Fig. 2. — A-C, Cheliplana pileola n. sp.; A, squeezed live specimen (free-hand drawing); B, proboscis; C, maie copulatory organ (B-C from whole mount). D-F, Cheliplana textilis n. sp.; D, squeezed live specimen (free-hand drawing); E, proboscis; F, male copulatory organ (E-F from whole mount). Kalyptorhynchia from Kenya 153

Material. Four animals studied alive, three mounted (holotype LUC No. 144, paratypes LUC No. 145 and 146) and one serially sectioned (paratype LUC No. 147). Description. The species is 720-860 pm long (measured on the fixed animals). Caudally, a haptic girdle and haptic glands are present. There are no pigmented eyes. The cylindrical pharynx (about 120 pm long) is situated .in the anterior third of the body. The long pharynx cavity bears no spines but cell bodies with nuclei protrude above llie surf are of the cavity epithelium (sectioned material). The curved proboscis hooks, without denticles, are 14-18 pm long. Soft side pieces without sclerotic rods are present as well. The muscle bulbs of the proboscis are 7.5-13 pm long. The unpaired, slightly bilobed testis, situated beside and behind the pharynx, is probably a product of fusion of paired testes. The male copulatory organ is located in the caudal quarter of the body. The paired seminal vesicles lead into the small prostatic vesicle (25-30 pm long), which contains only one type of prostatic secretion. The 45-50 pm long cirrus is enveloped in a 20 pm long sclerotic papilla at the distal end. Proximally, the spines are 2.5-4 pm long, distally 3-6.5 pm. Separately, two other gland systems empty into the male atrium, together with a blind diverticle with unknown function. The large ovary is located beside the cirrus. The yolk glands are paired and reach from behind the testis to the ovary. Behind the ovary, a large bursa is present. The presence or absence of an external vagina could not be ascertained with certainty. If an external vagina is present it is situated behind the common genital pore. Discussion. See general discussion for bothCheliplana- species.

Cheliplana textilis n. sp. (Figs 2D-F and 3I-J) Locality in Kenya. Same locality as forC. pileola and at English Point, in a seepage area at mid-tide level, where water of lower salinity (28%o) bubbled up (in very fine sand, mixed with some plant debris).

Material. Four animals studied alive, one mounted (holotype LUC No. 148) and three serially sectioned (paratypes LUC No. 149, 150 and 151). Description. The fixed animals are 300-480 pm long. A caudal haptic girdle is present. The cylindrical pharynx is connected to the mouth pore by a long pharynx cavity without spines. The proboscis is 20 pm long with 16 pm long hooks and short (6 pm) muscle bulbs. It has soft side pieces with small sclerotic needles. The unpaired testis is situated beside and in part behind the pharynx. The male copulatory organ lies in the posterior half of the body. Two seminal vesicles are present, the copulatory bulb is 50 pm long. Beside the two seminal vesicles, two strands of prostatic glands enter the bulb and surround the ejaculatory duct in the proximal part of the bulb. The copulatory bulb contains a 30 pm long cirrus, which is «entwined» by two additional blind cirri (each about 40 pm long). The distal part of the bulb is surrounded by a 17.5 pm wide cap. The ejaculatory cirrus is completely lined with small spines (1-3 pm long). The two additional cirri have only spines on one side of the cirrus, and in the posterior half. Beside these cirri, another group of prostatic glands are enclosed in the copulatory bulb. The large, unpaired ovary lies beside the large common genital atrium. The strong longitudinal muscles surrounding the genital atrium are continuous with the 154 P.E.H. Jouk and A.J.-P. De Vocht

20 firn

Fig. 3. — A,Polycystis ali, stylet; B, Alcha evelinae, male cuticular apparatus; C-D, Carcharodorhynchus involutus n. sp.; C, proboscis and D, male copulatory organ; E-F, Baltoplana bisphaera n. sp.: E, male copulatory organ: copulatory bulbus and vesiculae seminales and F, diverticles; G-H,Cheliplana pileola n. sp.: G, proboscis and H, male copulatory organ; I-J, Cheliplana textilis n. sp.: I, proboscis and H, cirrus. (All pictures taken from whole mounts). Kalyptorhynchia from Kenya 155

body wall musculature (sectioned material). The unpaired strand of yolk glands extends from just behind the pharynx to the ovary, A large bursa is found behind the ovary. Discussion. The generic position ofC. pileola and C. textilis approves to be indisputable. They have proboscis hooks without denticles and with soft side pieces, a cylindrical pharynx with a long cavity, one testis and one ovary and a male copulatory organ with eversible cirrus. The interspecific relationship of both species in the genus cannot fully be discussed, because our knowledge of the internal organisation is only fragmentary. C. pileola shows a resemblance toC. barringtonensis Nodt & Hoxhold19 8 4 and C. pacifica Noldt & Hoxhold1 9 8 4 , two species from the Galapagos Islands and C. californica Karling 1 9 8 9and C. elkhornica Karling 1 9 8 9from California. However, C. barringtonensis differs from C. pileola in having a spinous pharynx cavity and a much larger prostatic vesicle(1 2 0 pm). The ejaculatory duct of the former forms10-14 a pm long sclerotic tube which widens into a short cirrus(2 0 -2 5 pm) with uniform small spines(2-3 pm). The cirrus narrows distally into a fine channel, distally everted to a 12 pm long sclerotic papilla(Noldt & Hoxhold 1984). C. pacifica also has a cirrus which distally is enclosed in a sclerotic papilla. The cirrus5 0 -6is 0 pm long (spines1-2 pm), the sclerotic papilla2 5 -3 0 pm. Furthermore, it has a large prostatic vesicle(220- 2 7 0 pm) and a very long ejaculatory (Nduct oldt & H oxhold 1984). C. barringtonensis and C. pacifica have an unpaired strand of yolk glands. BothC. californica and C. elkhornica possess a relative short ejaculatory duct and cirrus, with sclerotic papilla. An external vagina is present in both species and situated in front of the common genital pore. C. textilis is the only species in this genus with two additional blind cirri beside the ejaculatory cirrus. C. asica Marcus 1 9 5 2 has a ductus ejaculatorius without spines.C. In asica terminalis the terminal part of the ejaculatory duct and the wall of the penial papilla are sclerotic (B r u n e t 1 9 6 8 , fig. 3). A sclerotic papilla formed by a variable everted part of the ejaculatory duct is present in many species of the genus (and other genera) and has only a restricted taxonomic value. The relative length of proboscis hooks and muscular bulbs might prove to be a valuable instrument to find interspecific relation­ ships within the genus.

ABBREVIATIONS

ac\ additional cirrus; b\ bursa; br. brain; bt: bursal tissue; ca: cuticular cap; cb: copulatory bulbus; ci\ cirrus; d: diverticle; gu gland of type 1; g2: gland of type 2; gg,: genital gland of type 1; gg2: genital gland of type 2;md: muscular duct; ov. ovary;p: genital pore; pg¡: prostatic gland of type 1; pg2: prostatic gland of type 2; ph: pharynx;phc: pharynx cavity;pr: proboscis; prg: proboscis glands; pi: proboscis teeth; s: stylet; sp', cuticular spine; t: testis; v. vitellaria; vs: vesicula seminalis.

ACKNOWLEDGEMENTS

The first author wishes to thank Prof. Dr P. Polk (Director of the Kenyan-Belgian Project in Marine Ecology) for the opportunity to come to Kenya. He also thanks Dr E. Martens (Assistant- Director of the project) for her help and assistance during the second stay. Thanks are also due to 156 P.E.H. Jouk and A.J.-P. De Vocht

Mr S.O. Alíela (Director of K.M.F.R.T.) for providing working facilities at the institute and to Mr G. Wamukoya (K.M.F.R.I.) for identification oi the algae. Both authors wanL Lo express their gratitude to Prof. Dr E. Schockaert, Dr P. Martens and Prof. Dr T. Karling for valuable discussions and comments.

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