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A New Genus and Species of Lineid Heteronemertean from South Africa, Polybrachiorhynchus Dayi (Nemertea: Anopla), Possessing a Multi- Branched Proboscis

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A New Genus and Species of Lineid Heteronemertean from South Africa, Polybrachiorhynchus Dayi (Nemertea: Anopla), Possessing a Multi- Branched Proboscis BULLETIN OF MARINE SCIENCE, 27(3): 552-571, 1977 A NEW GENUS AND SPECIES OF LINEID HETERONEMERTEAN FROM SOUTH AFRICA, POLYBRACHIORHYNCHUS DAYI (NEMERTEA: ANOPLA), POSSESSING A MULTI- BRANCHED PROBOSCIS Ray Gibson ABSTRACT A new genus and species of lineid heteronemertean from South Africa, Polybracllio- rhyne/Ius dayi, is described and illustrated. lts most characteristic taxonomic feature, the mode of branching of the proboscis, distinguishes the worm from all other described nemerteans. The proboscis consists of a single main axis from which the primary branches arise in only a single plane either alternately along the lateral margins or dichotomously at the distal tip. Each primary branch then bifurcates regularly up to the quaternary branch let stage. Other anatomical characters can also be used to distinguish the new taxon of worms from existing heteronemertean species which have a branched proboscis and these are discussed in detail. A key for the identification of nemertean species which possess a branched proboscis is provided. Only three species of nemerteans, all be- and Panorhynchus. They defined Gorgono- longing to the anoplan order Heteronemer- rhynchus as possessing a branched proboscis, tea, have so far been described with a the branches being regularly dichotomous branched proboscis. Gorgonorhynchus re- with numerous (32-64) divisions, whereas pens Dakin and Fordham (1931) has been the structure from Panorhynchus was de- reported from Australia (Dakin and Ford- scribed as forming a panicle-like branching ham, 1931, ] 936; Dakin, 1973; Bennett, system, with the proximal portion of the 1974) and the Marshall Islands (Coe, proboscis trunk containing one muscle cross 1947), and is probably the form from and a complete layer of circular musculature. India referred to by Gravely (1927) (see Further differences between the proboscides Gibson, 1974a: 473-474); Gorgonorhyn- of these genera lie in that "The characteristic chus bermudensis Wheeler (1940a) is dendritic form of the everted proboscis (of known from Bermuda (Wheeler, 1936, Gorgonorhynchus) is achieved by a regular 1940a) and Florida (Gibson, 1974a); and alternation of the planes of division" (Dakin Panorhynchus argentinensis Serna de Este- and Fordham, 1936: 465), whereas in ban and Moretto (1969) has been found Panorhynchus the principal lateral branches only in a lagoon near Buenos Aires, Argen- emerge in a "very open spiral" (Serna de tina (Serna de Esteban and Moretto, 1969). Esteban, pers. comm. 23 February 1976). Few species of nemerteans have been re- The proboscis of the new South African form corded from South African shores (Wheeler, possesses quite a different mode of branch- 1940b; Day, 1974). Among these, Day ing; the primary branches, apart from the (1974: 47) lists a "Gorgonorhynchus sp.," terminal pair which result from a simple although his illustration depicts a proboscis dichotomous division of the distal axial which is not branched dichotomously as in tip, are given off laterally and alternately the two species of this genus. Indeed, the from the single main axis entirely within a nature of the proboscis branching was used single plane. Each primary branch then by Serna de Esteban and Moretto (1969: divides through three successive bifurcations 167) as a major taxonomic character for to give eight quaternary branchlets. This distinguishing between Gorgonorhynchus manner of branching, together with other 552 GIBSON: NEW GENUS OF HETERONEMERTEAN 553 morphological differences, excludes the trichrome or Alcian blue methods for ana- present nemerteans from either of the genera tomical investigations. listed above and they are accordingly placed In addition, the morphology of the present in a new genus, Polybrachiorhynchus specimens has been compared with selected (Greek: poly-many; brachion-an arm; stained slides of Panorhynchus argentinensis rhynchos-a snout), the specific name dayi kindly loaned by Dr. Carmen J. de la Serna being chosen in honour of the finder, Pro- de Esteban, University of Buenos Aires. fessor John H. Day, and to mark his retire- ment from the Department of Zoology, Uni- Polybrachiorhynchus new genus versity of Cape Town, South Africa. Heteronemerteans with a single pair of deep horizontal lateral cephalic slits; with a MATERrALS AND METHODS complex branched proboscis comprising an The description of Polybrachiorhynchus undivided main axis from which primary dayi is based upon five specimens collected branches emerge in a single plane, terminal by Professor Day. Three of the animals, primaries formed by dichotomy of the distal originally given to the British Museum (Nat- axial tip but remaining primaries arising ural History), London, several years ago, alternately from the lateral axis margins, have been loaned through the courtesy of each primary branch dividing dichotomously Mr. S. Prudhoe but two were sent directly to to give eight quaternary branchlets; pro- the author by Dr. Charles Griffiths of the boscis axis containing three muscle layers Department of Zoology, University of Cape (outer and inner longitudinal, middle circu- Town. The localities from where the nemer- lar) and two muscle crosses; rhynchocoel teans were obtained, together with their musculature not interwoven with body wall University of Cape Town Ecological Survey muscle layers; dermal glandular zone and Catalogue Numbers, are: (1) BRE 73A outer longitudinal musculature of body wall Breede River. 34°S, 200E. One specimen, separated by distinct layer of connective tis- dug from mud in Zostera zone. Collection sue; dorsal fibrous core of cerebral ganglia date unknown. (2) BRE 158B Breede simple, not forked either at front or back; River. Two specimens, near river mouth, nervous system with neurochord cells in the intertidally from Upogebia beds. Collected 9 brain and neurochord fibres in the lateral January 1974. (3) DBN 54B Durban Bay. nerve cords; proboscis nerve supply a neural One specimen, burrowed in a sand bank. sheath without distinct nerves; foregut with Collected 19 July 1950. (4) DBN 211A a delicate layer of circular splanchnic Durban Bay. One specimen, swimming next muscles and with a longitudinal muscle plate to a wharf at night. Collected 4 October between dorsal wall and rhynchocoel; main 1951. intestinal canal enclosed by a layer of longi- The worms were preserved in formalin but tudinal muscle fibres; blood system with a have subsequently been secondarily fixed in cephalic lacunar network and a postcerebral hot (60°C) Bouin's fluid prior to sectioning foregut plexus, transverse blood connectives at 6-7 ""m in 56°C melting point paraffin present in intestinal region, mid-dorsal vessel wax. The smaller of the two worms holding developed into a long rhynchocoelic villus; catalogue number BRE 158B has been com- cephalic glands well developed, reaching pletely sectioned, but the large size of the backwards to anterior margins of brain and remaining animals made it impracticable to opening via frontal organ; without eyes; section them fully and accordingly only rep- with a caudal cirrus. resentative portions of their bodies have been examined histologically. All sections Type-Species.-Polybrachiorhynchus dayi, have been stained by either the Mallory new species. 554 BULLETIN OF MARINE SCIENCE, VOL. 27, NO.3, 1977 Polybrltchiorhynchus dltyi new species Type-Specimens.-Type material deposited with the British Museum (Natural History), London, consists of series of stained sec- tions and uncut body fragments. The Registration Numbers, with relevant Uni- versity of Cape Town Catalogue Numbers listed in brackets, are: Holotype, fully sec- tioned specimen, 1976.3.18.2 (BRE 158B); Paratypes, 1976.3.17.1 (DBN 211A), 1976.3.17.2 (DBN 54B), 1976.3.18.1 (BRE 73A) and unsectioned animal under 1976.3.18.2 (BRE 158B). Type-Locality.-Breede River, South Af- rica, 34°24'S, 20050'E, near river mouth, intertidally from Upogebia beds. External Appearance.-Living worms are coloured "straw yellow or tinged with pink" (letter from Professor Day dated 10 Jan- uary 1974) or yellowish-white (Day, 1974). Colour notes accompanying two of the speci- mens record that DBN 54B was reddish brown overall with pale edges, whilst DBN 211A possessed a pale yellow-brown head but had a darker body with transparent pink Figure 1. PolybrachiorhYllchus dayi. External margins. Preserved animals are a uniform features, drawn from preserved specimen. (A) En- dull flesh colour overall. tire worm. cc, caudal cirrus; If, flattened lateral 'fins' of intestinal region. (B) Anterior end viewed Polybrachiorhynchus dayi attains a size from ventral surface. mt, mouth; pp, proboscis considerably larger than has been reported pore. (C) Anterior end from lateral aspect. cs, for either Gorgonorhynchus or Panorhyn- cephalic slit. The scale bar refers to A only. chus species; live individuals, after anaesthe- tization, reach a maximum of some 25 in. in Distinct "fins" extend along the lateral mar- length (ca. 60 cm) (letter from Dr. Grif- gins for most of the body length, much as in fiths, 29 January 1974), compared with other nemerteans (e.g., species of Cerebratu- about 20 cm for Gorgonorhynchus repens Ius) which are capable of actively swimming (Dakin and Fordham, 1936), 12 cm for in an eel-like manner (Verrill, 1892; Coe, Gorgonorhynchus bermudensis (Wheeler, 1943). Posteriorly the body gradually tapers L940a) and 18 cm for Panorhynchus argen- to terminate in a small stumpy caudal cirrus tinensis (Serna de Esteban and Moretto, 3 mm or less in length. The wedge-shaped 1969). The body is
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