\3 Biodiversity ^Heritage ^^Library http://www.biodiversitylibrary.org/ Proceedings of the Biological Society of Washington. Washington, Biological Society of Washington http://www.biodiversitylibrary.org/bibliography/3622 v. 102 Mar-June 1989: http://www.biodiversitylibrary.org/item/107602 Page(s): Page 1 1 6, Page 1 1 7, Page 1 1 8, Page 1 1 9, Page 1 20, Page 1 21 , Page 1 22, Page 1 23 Contributed by: Smithsonian Institution Libraries Sponsored by: Biodiversity Heritage Library Generated 19 November 2011 9:30 PM http://www.biodiversitylibrary.org/pdf3/008713500107602 This page intentionally left blank. PROC. BIOL. SOC. WASH. 102(1), 1989, pp. 116-123 REVISION OF THE FAMILY LYSARETIDAE, AND RECOGNITION OF THE FAMILY OENONIDAE KINBERG, 1865 (EUNICIDA: POLYCHAETA) G. Kent Colbath Abstract. — As recognized by previous authors, the family Lysaretidae Kin- berg, 1865 includes genera with dissimilar jaw morphology and composition. Lysaretidae is restricted here to include only Lysarete Kinberg, characterized by jaws mineralized with calcite and relatively short carriers with transverse musculature. Oenone Savigny, Halla Costa and Tainokia Knox & Green are removed into the family Oenonidae Kinberg, 1865. These genera are charac- terized by heavily sclerotized jaws which are not mineralized, and by long carriers with oblique -longitudinal musculature. Hartman (1944) published a family-level revision of the eunicoid polychaetes which was widely accepted. Drawing on the work of Kinberg (1865) and Ehlers (1868), she placed heavy emphasis on the configuration of the jaws as a taxonomic character. Hartman recognized three basic maxil- lary jaw patterns within the superfamily Eu- nicea: the labidognath pattern ascribed to the Eunicidae, Onuphidae, and Lumbriner- idae; the prionognath pattern in the Arabellidae and Lysaretidae; and a third pattern (later named ctenognath by Kielan- Jaworowska 1966) found only in the Dor- villeidae. Labidognath jaws are character- ized by having two relatively short, broad carriers, in contrast to the two long, slender carriers with a ventral median ligament (or "third carrier") which characterize priono- gnath jaw apparatuses. In the ctenognath condition the maxillae consist of two to four rows of numerous separate denticles. During a study of the composition and fossilization potential of polychaete jaws, I divided eunicoid jaws into three groups based on their composition (Colbath 1986). Labidognath jaws are mineralized with cal- cium carbonate in the form of aragonite in the Onuphidae and Eunicidae, and calcite in the Lumbrineridae. Labidognath jaws are also weakly sclerotized, and the outer pro- tein wall is readily leached in 2% KOH (Hartmann-Schroder 1967, Colbath 1986). In contrast, prionognath and ctenognath jaws are amorphous to x-rays, do not react with HC1, and are relatively resistant to leaching in KOH (Colbath 1986). The jaws of species of Lysarete Kinberg, 1865 are mineralized with calcite (Colbath 1986), and exhibit other labidognath mor- phological characters. Other genera previ- ously included within the Lysaretidae have jaws of the prionognath type, prompting a revision of the family, and recognition of the family Oenonidae Kinberg, 1865. History of Study Early classifications of the eunicoid poly- chaetes were based exclusively on features of the external anatomy, and produced ar- bitrary groupings of genera which are dis- similar internally (Hartman 1944:1). First to emphasize the importance of jaw parts in classification was Kinberg (1865). Kin- berg recognized ten families which he di- vided into four major groups (labeled in- formally as A-D). He defined group C by the presence of mandibles with separated right and left halves (an ontogenetically variable feature), and included only La- VOLUME 102, NUMBER 1 117 randa Kinberg, 1865, which was considered unrecognizable by Hartman ( 1 944: 1 0, 1 948: 98). Kinberg's other groups correspond to the labidognath (A), prionognath (B), and ctenognath (D) jaw types presently recog- nized. Kinberg considered the Lysaretae (sic) and Oenonidea (sic) distinct, including the former in group A and the latter in group B. Ehlers (1868:280-282) also emphasized the importance of jaw configuration in clas- sifying these polychaetes, but did not for- malize the distinctions. Instead, he recog- nized a single family Eunicea (sic), which he informally divided into the Eunicea lab- idognatha and Eunicea prionognatha. Eh- lers (1868) did not stress the morphology of maxillary carriers as strongly as did Kin- berg, but rather subdivided the eunicoids according to whether the anterior maxillae are arranged in a semicircle (labidognatha) or in parallel rows (prionognatha). In contrast to Kinberg, Ehlers (1868) placed the genus Lysarete close to Aglau- rides Ehlers (=Oenone Savigny) in the prionognath subdivision. He later (Ehlers 1887:107-108) acknowledged the close similarity between the jaws of Lysarete and those of Lumbriconereis Grube (=Lumbri- neris Blainville), but did not emend his ear- lier taxonomic treatment. Hartman (1944:2) essentially formalized the 1 868 scheme of Ehlers using the family- level nomenclature of Kinberg. She retained six of the ten families recognized by Kin- berg, placing the Oenonidae into synonymy with the Lysaretidae. Fauchald (1970:118) removed Iphitime Marenzeller from the Lysaretidae, and erected the family Iphitimidae. He later (Fauchald 1977) erected the order Eunicida to include the seven families within the su- perfamily Eunicea, plus two additional fam- ilies (Histriobdellidae, Ichthyotomidae) which were not assigned to a superfamily. Materials and Methods All lysaretids in the collection of the U.S. Museum of Natural History (USNM) were examined for the present study, and selected specimens were dissected for examination of the jaws. Additional specimens of Ly- sarete were obtained on loan from the Flor- ida Department of Natural Resources (FSBC). The holotype of L. brasiliensis Kin- berg, 1865 was obtained from the Swedish National Museum, Stockholm. One specimen of Lysarete brasiliensis with jaws in excellent condition (USNM 55884) was used for the chemical and mineralogical analyses, and for scanning electron micros- copy. The maxillary apparatus was re- moved, dehydrated in an alcohol series, and prepared for SEM by critical point drying. The specimen was initially coated with car- bon only, and chemically analyzed using x-ray dispersion. The specimen was then coated with gold palladium for routine scan- ning electron microscopy. The mandibles were removed, photographed with a Wild 400 wide-field optical microscope, and ana- lyzed with the Gandalfi x-ray spectrometer in the Department of Mineralogy, Smith- sonian Institution. Taxonomic Characters Jaw morphology and composition. —X-ray dispersion analysis of the maxillae of Ly- sarete brasiliensis indicates the presence of abundant calcium carbonate. The mandi- bles are mineralized with calcite, as are jaws in the lumbrinerids (Colbath 1986). Calci- fication is also evident in L. australiensis Benham, 1915, in that the blades of the mandibles are white, and the maxillae are dark brown rather than black. The presence of calcite is inferred by comparison with L. brasiliensis. In contrast to Lysarete, jaws in species of Oenone Savigny, Halla Costa and Tainokia Knox & Green (commonly included in the Lysaretidae) are not mineralized, but rather are composed almost exclusively of sclero- protein. As a consequence, both maxillae and mandibles are jet black in color. The distinction between heavily sclerotized and calcified jaws is apparent upon inspection, 118 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON and was described by Benham (1915:233, 236-237). Benham made no comment about the utility of this distinction above the species level, however. Other aspects of jaw morphology are cor- related with the difference in mineraliza- tion. In Lysarete the maxillary carriers are relatively short (approximately equal to the length of the maxillae) and broad (Fig. lb), with no ventral ligament (= "third carrier"). A thick band of transverse muscles joins the carriers dorsally (Fig. 2f ), allowing them to be pulled closed in the "trap" motion de- scribed by Wolf (1980) for other labido- gnath jaws. The MI (maxilla I) elements in the maxillary apparatus are the same length (Fig. 2c), which allows the anterior tips to come together in a pincer-like fashion as part of the trap motion. The posterior dorsal surfaces of the Mis exhibit an interlocking ridge and furrow system (Fig. 2e), which may help to lock the apparatus in a closed position (Wolf 1980). In species of Oenone, Halla and Tainokia the carriers are narrow and much longer than the combined length of the maxillae, and are not equiped with a transverse muscle band. Instead, a series of long, oblique-lon- gitudinal muscles are attached along the length of the carriers. A ventral ligament is present. The right and left MI elements may be of different lengths (Crossland 1924; Hartman 1944; Kielan-Jaworowska 1966), and no basal ridge and furrow system is present to lock the jaws in a closed position. Such jaws are presumably incapable of per- forming a trap motion, as Wolf (1980) also concluded for the arabellids. Soft-part morphology. —The genera tra- ditionally included within the Lysaretidae share an elongated post-setal lobe and prominent dorsal cirrus on posterior para- podia. The morphology of the head, how- ever, is distinctly different in each of the four genera under discussion. Although the difference between Lysarete and the other genera may not merit consideration as a family-level character, the anterior mor- phology is none-the-less a useful key char- acter. In Lysarete, three antennae emerge from the posterior margin of the prostomium, and may lie flat in a V-shaped notch (Fig. la). In adult specimens the notch extends pos- teriorly across the peristomial rings, and in some cases across the first one or two seti- gers. Halla has three antennae which may lie back in a semicircular notch in the two peristomial rings (Fig. le). Tainokia has a single antennae, and a slight invagination which extends posteriorly only partially across the first of two peristomial rings (Knox & Green 1972, fig. 1). In Oenone, three antennae are tucked into a nuchal fold (Fig. Id), no notch is present, and dorsally the peristomium has only one distinct ring. Setae and acicula.— All of the worms considered here have simple geniculate se- tae. In addition, bifid hooks have been re- ported in Oenone and Halla. In Lysarete, Oenone, and Tainokia the geniculate setae are smooth (Knox & Green 1972, figs. 6, 7; pers. observ.). In Halla, however, the dorsal setae are denticulate, and the ventral setae are marked by distinctive oblique striae (Fig. 2a, b; Fauvel 1923:427, fig. 169 g, h). Both described species of Lysarete have black acicula. In Oenone fidgida (Savigny, 1818) and Halla parthenopeia (Chiaje, 1828) the acicula are amber, while Knox & Green (1972:433) described the acicula in Taino- kia iridescens as colorless. Black acicula are thus useful for distinguishing Lysarete from the other three genera, supporting the fam- ily-level distinction based on jaw morphol- ogy. Systematic s Order Eunicida Fauchald, 1977 Family Lysaretidae Kinberg, 1865 Emended diagnosis, — Eunicoid poly- chaetes with jaws mineralized with calcite. Mandibles flat with semicircular growth rings anteriorly (Fig. 2d). Maxillary carriers no longer than combined length of maxillae, VOLUME 102, NUMBER 1 119 e Fig. 1 . a-c, Lysarete brasiliensis, holotype, Brazil; a, Dorsal view, anterior end, illustrating V-shaped notch in peristomium; b. Dorsal view, MI elements and carriers, muscle attachment stippled; c, Labial view, MIIL- MVL. d, Oenonefulgida, USNM 17729, Dry Tortugas, dorsal view, anterior end, note tips of antennae protruding from nuchal fold, e, Hatta parthenopeia, USNM 5 147, Bay of Naples, dorsal view, anterior end, note semicircular notch in peristomium. All scale bars = 1 mm. articulated by prominent transverse mus- cular bundle, no ventral ligament. Maxillary jaws symmetrical. MI elements of identical length, distally falcate with prominent sec- ondary fang, additional small denticles may be present at base. Three short antennae arise from base of prostomium. Elongate post-setal lobe and flattened dorsal cirrus (or notopodium) developed in posterior parapodia. Simple geniculate setae present, bifid hooks absent. Comparison.— The Lysaretidae are dis- tinguished from the Oenonidae based on jaw morphology and mineralogy. Lysarete, the only genus presently recognized within the family, is further distinguished from the Oenonidae in having black acicula, and a V-shaped notch in the dorsal surface of the peristomium. The Lysaretidae are similar to the Lum- brineridae in having: 1) jaws mineralized with calcite; 2) flat mandibles with a semi- circular blade; 3) symmetrical maxillary jaws; 4) transverse musculature on the car- riers. The Lysaretidae are distinguished from the Lumbrineridae by: 1) well-developed 120 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. a, b, Halla panhenopeia, USNM 5147, Bay of Naples; a, SEM dentate setae from dorsal bundle, posterior parapodium, scale bar =10 fim; b, SEM striate setae from ventral bundle, scale bar = 50 (im. c-g, Lysarete brasiliensis, USNM 55884, Gulf of Mexico; c, SEM maxillary apparatus, dorsal view, scale bar = 1 mm; d, Light photomicrograph mandibles, ventral view, scale bar = 1 mm; e, SEM MI elements, oblique-lateral view, note ridges and grooves on bases of Mis, scale bar = 0.5 mm; f, SEM carriers, oblique-lateral view, carriers canted at an angle to Mis, note transverse muscle band, scale bar = 0.2 mm; g, SEM anterior maxillary jaws, oblique-lateral view, scale bar = 0.2 mm. dorsal cirri; 2) a prominent secondary fang on the MI elements of the maxillae; 3) the absence of hooded hooks. Kuwaita Moham- mad, 1973 is intermediate between the Ly- saretidae and Lumbrineridae in having an- tennae on the prostomium, but clearly belongs in the Lumbrineridae based on the 3 criteria above (Mohammad 1973:34, 36, figs. 6, 7). Distribution.— Lysarete includes one VOLUME 102, NUMBER 1 121 species recorded from the east coast of North and South America in low latitudes, and a second species from a single locality east of Australia. Larymna Kinberg, 1865 may also belong in the Lysaretidae based on jaw mor- phology (Hartman 1948:99). The type and only specimen (from Mozambique) is so poorly preserved that, according to Hart- man, attribution cannot be made below the family level, and Larymna should be re- garded as unrecognizable. Genus Lysarete Kinberg, 1865 Type species.— L. brasiliensis Kinberg, 1865:570. Lysarete brasiliensis Kinberg, 1865 Figs, la-c, 2c-g Lysarete brasiliensis Kinberg 1865:570; 1910:49, pi. 17,fig.30.-Ehlers 1887:107- 108, pi. 33, figs. 1-8.— Rioja 1944:130- 131. figs. 47-50.— Hartman 1951:65, pi. 14, figs. 7, 8.-Orensanz 1975:106-108, fig. 8. -Gardiner 1976:213, figs. 29b-e.- Gilbert 1984:43-6-43-7, figs. 43-3, 43-4. lOenone brevimaxillata. — Treadwell 1931: 1-3, figs. 4-8. Remarks. — In her re-appraisal of Kin- berg's type material, Hartman (1948:8) did not consider the holotype of Lysarete bra- siliensis, but instead referred to an earlier revision by Ehlers (1887:107-108, pi. 33). Ehlers did not figure the Brazilian type of L. brasiliensis, but rather figured a specimen collected from Florida. The posthumous publication of Kinberg's plates (Kinberg 1910) did not include illustrations of the jaws of the holotype. The maxillae are there- fore illustrated here (Fig. lb, c). On the holotype, maxillary jaws III and IV each have a prominent anterior denticle, with two progressively smaller denticles posteriorly. The jaws in a specimen from Argentina (Rioja 1944, fig. 50) are similar, except that the Mill elements have four denticles instead of three. Most specimens examined from the east coast of the United States have five to six denticles on the Mill and four to seven den- ticles on the MIV elements (Fig. 2g). The anterior three denticles on the Mill element are sub-equal in length, unlike those of the type. One specimen from Florida (FSBC I 31008) has Mill jaws like the type, how- ever, and a specimen from Argentina illus- trated by Orensanz( 1975, fig. 8-3) has Mills and IVs indistinguishable from most of the North American specimens. There is thus some overlap in jaw morphology between the North and South American populations. The number and shape of denticles on an- terior jaws is variable in eunicoid poly- chaetes, and the two populations are ten- tatively considered conspecific in spite of their disjunct geographic ranges. The type and only specimen of Oenone brevimaxillata Treadwell, 1931 was never deposited (H. S. Feinberg, AMNH. pers. comm. 1985), and is presumed lost. Tread- well's (1931, fig. 8) illustration of the max- illae is schematic, and insufficient for de- tailed comparison. Family Oenonidae Kinberg, 1865 Emended diagnosis. —Jaws jet black, composed primarily of scleroprotein, no crystalline mineral phases present. Mandi- bles H-shaped. Maxillary carriers much longer than anterior jaws, attached to bun- dles of oblique-longitudinal muscles, ven- tral ligament present. Maxillae commonly asymmetrical, Mis differ slightly to mark- edly in length. One or three antennae arise from base of prostomium. Elongate post- setal lobe and prominent dorsal cirri de- veloped on posterior parapodia. Geniculate setae present, bifid hooks present or absent. Key to the genera of Oenonidae la. One or 3 antennae recline in notch in peristomium, 2 peristomial rings dorsally, bifid hooks present or ab- sent 2 1 b. Three antennae tucked into nuchal 122 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON fold, 1 peristomial ring dorsally, bifid hooks present Oenone 2a. One antenna, peristomial notch weakly developed, bifid hooks ab- sent, setae smooth Tainokia 2b. Three antennae, distinct semicir- cular peristomial notch, bifid hooks present (small individuals only), dorsal setae dentate, ventral setae Genus Halla A. Costa, 1844 Type species. —Halla parthenopeia (Chiaje) Costa, 1844:63.— Lysidice parthe- nopeia Chiaj e, 1828:175. Remarks. — Fauvel (1923:427, fig. 169f) described Halla parthenopeia as having bi- fid hooks, in addition to the setae figured here (Fig. 2a, b). Hooks are absent in para- podia from the large specimen of H. par- thenopeia examined here (USNM 5 147), nor were hooks recorded from the large speci- men figured by Ehlers (1868:28, figs. 27, 28). The type of Halla okudai Imajima (1967:437) is also a large worm, and lacks hooks (Imajima & Hartman 1964:267-268). Individuals of Halla which attain large size may lose their bifid hooks. A similar con- dition prevails in several large species with- in the Eunicidae (K. Fauchald, pers. comm. 1985). Distribution. — All western hemisphere specimens deposited in the USNM collec- tion as Halla proved to be misidentified members of Oenone or Lysarete. Halla may be restricted to the Old World, with docu- mented reports from the Mediterranean (Fauvel 1923) and Japan (Imajima 1967). Acknowledgments I thank Kristian Fauchald for extensive discussions about the taxonomy of eunicoid polychaetes, and for critically reading the manuscript. Thomas H. Perkins arranged the loan of material from the Florida De- partment of Natural Resources. H. S. Fein- berg, American Museum of Natural His- tory, provided information concerning the type of Oenone brevimaxillata Treadwell. This research was supported by a post-doc- toral fellowship, Smithsonian Institution. Literature Cited Benham, W. B. 1915. Report on the Polychaeta ob- tained by the F. I. S. "Endeavour" on the coasts of New South Wales, Victoria, Tasmania and South Australia, part I. — Biological Results of the Fishing Experiments carried on by the F. I. 5. "Endeavour," 1909-14:173-273. Chiaje, S. delle. 1 828. Memorie sulla storia a notomia degli animali senza vertebre del regno di Napoli, 3. Napoli, 232 pp. Colbath, G. K. 1986. Jaw mineralogy in eunicean polychaetes (Annelida),— Micropaleontology 32: 186-189. Costa, A, 1 844, Cerai intorno alle osservazioni zool- ogiche fatte durante intre mesi vernali del 1844.— Annales Accademia aspirantes Natu- rales Napoli 2:40-68. 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