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Riserins Pugetensis Gen. N., Sp. N. (Nemertina: Anopla), a New Mesopsammic Species, and Comments on Phylogenetics of Some Anoplan Characters

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Riserins Pugetensis Gen. N., Sp. N. (Nemertina: Anopla), a New Mesopsammic Species, and Comments on Phylogenetics of Some Anoplan Characters Hydrobioiogia 266: 203-205, 1993. R, Gibson, J. Moore & P. Sundberg (eds). Advances in Nemenean Biology 203 © 1993 Kluwer Academic Publishers. Printed in Belgium. Riserins pugetensis gen. n., sp. n. (Nemertina: Anopla), a new mesopsammic species, and comments on phylogenetics of some anoplan characters Jon L. Norenburg Department of Invertebrate Zoology, National Museum of Natural History, Washington, DC 20560, USA Key words: nemertine, systematic s, phylogeny, meiofauna Abstract Riserius pugetensis gen, n., sp. n. is described from the northwest coast of Washington, U.S.A. It is probably a heteronemertine and, as such, would be the first known mesopsammic member of that order; it lives in the interstices of medium to coarse, moderately clean sands. Its morphology presents some attributes considered characteristic of mesopsammic fauna. The effect of some of these attributes is a generalized appearance of anatomical 'simplicity' and, as with other mesopsammic metazoans, this presents difficulties in distinguishing between primitiveness and reduction. In the new species such problematic features include the lack of subepidermal glandular cells and connective tissue, reduced proboscideal musculature, simple blood-vascular system, and the presence of a 'secondary', outer cir- cular musculature in the body wall. The general appearance of this new species is very similar to the so-called palaeonemertine Hubrechtella and characteristics of both suggest relationship with the baseo- discid heteronemertines. These relationships are explored in light of a modified interpretation of pro- boscideal musculature, a traditional cornerstone of heteronemertinean taxonomy. Introduction (Gibson, 1982). Palaeonemertina (sensu Iwata, 1960) are Anopla having the nervous system at Riseliuspugetensis gen. n., sp.n. fits a basic struc- the base of the epidermis and lacking outer lon- tural definition of the Heteronemertina but re- gitudinal musculature and subepidermal gland sembles palaeonemertines in its simplicity. The cells. Although not explicit on the issue, Burger orders Heteronemertina and Palaeonemertina are (1895) effectively recognized the palaeonemer- usually defined on the basis of body wall organi- tines as paraphyletic, but the heteronemertines as zation, arrangement of the dermis, and position monophyletic. In contrast, Hylbom's (1957) phy- of the nervous system (e.g, Hylbom, 1957; Gib- logenetic scheme, which echoed the opinions of son, 1982). Heteronemertina are defined as hav- several other principal nemertinologists, results in ing a body wall constructed as follows: an outer both orders being at least paraphyletic. A 'cuds' zone (dermis) containing connective tissue, sub- comprising the subepidermal cell bodies of glan- epidermal glands and sometimes circular muscle dular cells and often an associated zone of con- fibers; an outer longitudinal musculature; the cen- nective tissue, is characteristic of heteronemer- tral nervous system between this longitudinal tines. Potential homologues of these cutis musculature and a deeper circular musculature glandular cells are known from several palaeone- 204 mertinean genera, e.g., Hubrechtelh Bergendal, a 63-ftm mesh sieve. The worms were examined 1902, in which submerged cell bodies form a re- while alive with a stereomicro scope and, while gion of pseudostratified epidermis (Gibson, lightly squeezed under a coverslip, with a com- 1979a; Norenburg, 1985). Riserius pugetensis has pound microscope utilizing bright-field or Nomar- neither pseudostratified epidermis nor compo- ski optics. For preservation and histology speci- nents of a cutis, a condition known among other mens were anaesthetized briefly (5-6 min) in heteronemertines only from the lineid Coleman- 7.5% MgCl2, fixed in Hollanders cupri-picri- nietta albulus (Gibson, 1981). formal-acetic fixative, and embedded in polyester Effectively, the order Heteronemertina can be wax (Steedman, 1960; Norenburg & Barrett, separated from the order Palaeonemertina (sensu 1987). Transverse and longitudinal serial sections Iwata, 1960) only by the presence of the outer were cut at 5-7 fim and stained with Heidenhain's longitudinal musculature lying between the epi- azan or Heidenhain's iron hematoxylin methods. dermis and the central nervous system; i.e., this is a synapomorphy for Heteronemertina. How- Systematic account ever, less discrete longitudinal musculature may also be found in the dermis or among the epider- Riserius pugetensis gen.n., sp.n. mal cells of Palaeo- and Hoplonemertina (Friedrich, 1936, 1979). Similarly, some species Generic diagnosis of palaeo- and heteronemertines have a more or Four body-wall muscle strata, outer circular, less distinct dermal or epidermal outer circular outer longitudinal, middle circular, and inner lon- musculature, as seen in R. pugetensis. gitudinal; cutis (subepidermal glandular cells and Riserius pugetensis has been alluded to previ- connective tissue) lacking. Central nervous sys- ously as a mesopsammic heteronemertine tem and plexus between outer longitudinal and (Norenburg, 1987, 1988a, b). However, one may middle circular musculature; dorsal ganglia not legitimately ask, is it in fact a heteronemertine bifurcated posteriorly; without neurochord ele- and, if so, is it a relict species of a largely extinct ments; middorsal nerve from dorsal commissure. clade or is its simplicity a derived function of its Proboscis with inner longitudinal musculature; specialized habitat? inner and outer circular musculatures reduced or lacking. Rhynchocoel short; with outer circular Material and methods and inner longitudinal musculature. Cerebral or- gans not fused to cerebral ganglia; not projecting Six specimens of Riserius pugetensis gen. n., sp. n. into vascular lacunae; open via lateral pits. were collected in August, 1979, from sand at 0 m Transverse cephalic furrow encircles body in front (near the bottom of a moderate slope), 200 m of mouth. Rhynchodaeal aperture terminal; south of an oil terminal and depot at Richmond mouth ventral, posterior to cerebral organs. Blood Beach along Puget Sound, Washington vascular system with single cephalic lacuna; two (47° 40' N, 122° 25' W). Three more specimens lateral vessels and one middorsal; supra-anal were obtained from sand samples collected by anastomosis. Cyrtocytes single and clustered, divers in July, 1981, from Minnesota Reef near projecting into lateral vascular lacunae; one pair Friday Harbor, San Juan Island, Washington of collecting ducts and nephric pores. (48°31'48"N, 122°58'55"), at about 10 to 15 m depth. Sand samples were collected with a trowel Riserius pugetensis gen.n., sp.n. to a depth of 5-10 cm, placed in buckets or plas- tic Whirl-Paks and brought back to the labora- Specific diagnosis tory. Specimens were separated from the sand by Characters of the genus. Cephalic glands and swirling a subsample of sand in a bucket with frontal organ lacking. Inner circular musculature seawater and quickly decanting the water through reduced and longitudinal muscle plate lacking. 205 Blood vessels simple, lacking cross-connections, eggs are somewhat stouter. The epidermis is middorsal vessel enters rhynchocoel, precerebral translucent white with transmitted light and lacuna a single, medial sac. Cerebral organs large; opaque white with reflected light. The body is openings deep lateral pits. Rhynchodaeum thin- moderately flattened dorsoventrally when gliding walled. Rhynchocoel approximately one-quarter but the cephalic region is cylindrical and has a of body length. Stomach and intestine histologj- cephalothricid appearance (Fig. 1). The cephalic cally distinct, without constrictions or diverticula. region is relatively elongate; in a 10 mm speci- Gonochoric. men, the pair of conspicuous pits leading to the cerebral organs is about 500 fim from the cephalic Type specimens tip. An epidermal furrow encircles the body Holotype (USNM 149925), and single paratype 500 /im farther posteriad, but anterior to the (USNM 149926) deposited in the Division of mouth (Figs. 1-3). This furrow is directed poste- Worms, National Museum of Natural History riad dorsally and anteriad ventrally. The cepha- (USNM), Smithsonian Institution, Washington, lic tip bears a small dimple, the rhynchodaeal DC; both specimens were collected 17 July pore, which is surrounded by a few elongate cilia, 1981, anaesthetized in magnesium chloride, fixed but these are shorter than typical sensory cilia. in Hollanders cupri-picri-formal-acetic fixative With gentle squeezing of living specimens under and subsequently transferred to 70% ethyl alco- a coverslip one can occasionally observe the pair hol. of dorsolateral nephric pores and also individual mucoid strings emanating from large gland cells. Additional specimens These appear to account for the sticky, multi- Two sets of serial sections of specimens, from the point haptic reaction exhibited by this species. Richmond Beach site, in the author's possession When severely agitated, members of this species will be deposited in the USNM at a later date. have a tendency to coil. There is no indication of One is a set of three slides bearing transverse a caudal cirrus. serial sections (stained with Heidenhain's iron hematoxylin) of the anterior one-fifth of a speci- men. The other consists of eight slides bearing Body wall transverse serial sections of two specimens side Epidermis in sections is 6-10/im thick, with a by side (stained with Heidenhain's azan or ciliary border of 4-5 ^m. Two cell types predomi- Heidenhain's iron hematoxylin). nate
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