Redescription of Echinoderes dujardinii (Kinorhyncha) with Descriptions of Closely Related Species
ROBERT R. HIGGINS
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S. Dillon Ripley Secretary Smithsonian Institution SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 248
Redescription of Echinoderes dujardinii (Kinorhyncha) with Descriptions of Closely Related Species
Robert P. Higgins
SMITHSONIAN INSTITUTION PRESS City of Washington 1977 ABSTRACT Higgins, Robert P. Redescription of Echinoderes dujardinii (Kinorhyncha) with Descriptions of Closely Related Species. Smithsonian Contributions to Zoology, number 248, 26 pages, 31 figures, 2 tables, 1977.—A key to the adults of the genus Echinoderes is followed by a redescription of E. dujardinii from both adult and juvenile specimens. North American records for E. dujardinii are corrected by the description of E. kozloffi, new species. Echinoderes pennaki, a species sympatric with E. kozloffi, is redescribed, and the latter's sibling species, E. pacificus, is discussed. Synonomies and distribution records are given for species mentioned.
OFFICIAL PUBUCATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: The coral Montastrea cavernosa (Linnaeus).
Library of Congress Cataloging in Publication Data Higgins, Robert P. Redescription of Echinoderes dujardinii (Kinorhyncha) with descriptions of closely related species. (Smithsonian contributions to zoology ; no. 248) Bibliography: p. Supt. of Docs, no.: SI 1.27:248 I. Echinoderes dujardinii. 2. Echinoderes. 3. Kinorhyncha—Classification. I. Title: Rede- scription of Echinoderes dujardinii (Kinorhyncha) .... II. Series: Smithsonian Institution. Smithsonian contributions to zoology ; no. 248. QL1.S54 no. 248 [QL391.A2] 591'.08s [595M83] 76-608300 Contents
Page Introduction 1 Methods 2 Acknowledgments 3 Key to Adults of the Genus Echinoderes 3 Echinoderes dujardinii Claparede 4 Echinoderes kozloffi, new species 13 Echinoderes pennaki Higgins 18 Echinoderes pacificus Schmidt 21 Echinoderes brevicaudattis, substitute name for E. brevispinosus Higgins .. 21 Literature Cited 24
Redescription of Echinoderes dujardinii (Kinorhyncha) with Descriptions of Closely Related Species
Robert P. Higgins
Introduction of Claparede's publication, also described this spe- cies from the same locality, naming it Echinodera The first published observation of a kinorhynch dujardini. is that of the French naturalist F£lix Dujardin These descriptions suffered from the authors' lack (1851), who based his account on specimens found of experience with the taxon, as have many sub- while examining material washed from algae col- sequent ones, including my own. Despite Zelinka's lected near St. Malo on the coast of Normandy in 1841. In a manner not uncommon throughout the (1928) more comprehensive description of E. du- history of the study of meiofauna, Dujardin was jardinii in his Monographic der Echinodera, cer- unable to place the new invertebrate in an existing tain taxonomic ambiguities have persisted. The higher taxonomic category although he recognized lack of preserved specimens, especially types, has its morphological affinities with acanthocephalans, compounded the problem; consequently, the re- rotifers, sipunculids, nematodes, tardigrades, and ported distribution of E. dujardinii, as well as various crustaceans ("copepods without legs"). Ten other species, must be considered with due caution. years after his discovery, Dujardin (1851) intro- Since it was described, E. dujardinii has been re- duced the first kinorhynch as "a tiny marine ani- ported from 26 localities including the northern mal, I'Echinodere, constituting an intermediate and southern coasts of Europe, the Black Sea, form between the crustaceans and worms." A few Canary Islands, Japan, and the northwestern coast years later Leuckart (1854) noted that he had seen of the United States. "I'Echinodere" at Helgoland in 1846 but had as- No specimens are available to confirm Chit- sumed that it was a dipteran larva. wood's (1964a) report of E. dujardinii from Tomales "L'Echinodere" became Echinoderes dujardinii Bay, California, one of the two northwestern U.S. Claparede, 1863 when the latter author described localities. Neither an unpublished photograph by specimens from St. Vaast la Hougue, not far from Chitwood (pers. comm.) of one of the specimens, St. Malo where Dujardin made his discovery. The nor sketches of specimens collected at the same following year, Gosse (1864), apparently unaware locality and time (1960) by Dr. Tor G. Karling (pers. comm.), indicate that the species from Tomales Bay is E. dujardinii. Specimens sent to me Robert P. Higgins, Smithsonian Oceanographic Sorting Cen- ter, National Museum of Natural History, Smithsonian Insti- from San Juan Island, Washington, by Dr. Eugene tution, Washington, D.C. 20560. N. Kozloff, resembled E. dujardinii sufficiently to
1 2 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY justify the tentative use of this name (Kozloft, partially overcome by reducing, by about 30 per- 1972; Merriman and Corwin, 1973); however, both cent, the amount of chloral hydrate used in the Kozloft and I expressed doubts that could be an- medium and by sealing the preparation with Mur- swered only by comparing the San Juan Island rayite soon after the fluid has solidified. In this specimens with specimens from either the type- series of preparations, some specimens were re- locality or localities (Naples and Trieste) where mounted in glycerin once they had been oriented Zelinka (1928) obtained E. dujardinii from his dorsoventrally. This is a procedure that should be redescription. My collection from the latter two used only when an adequate series of specimens are localities failed to provide specimens of E. dujar- available, since in transferring them many dorsal dinii but Kozloff, in 1973, was successful in collect- spines are often broken. A few specimens were ing this species at Roscoff, not far from St. Vaast mounted in glycerin without first mounting them la Hougue. in Hoyer's medium. In all instances, coverslips were The purpose of this paper is to redescribe E. du- sealed with Murrayite. jardinii, clarifying both the original description by Each specimen was studied with the use of Zeiss Glaparede (1863) and the redescription by Zelinka differential interference contrast optics and me- (1928). In addition, this paper will correct the dis- ristically analyzed. The resulting data are expressed tribution record inasmuch as the San Juan Island in a standard format of abbreviations and termi- specimens, reported as E. dujardinii, constitute a nology (Higgins, 1967, 1969a). Measurements are new species described below. Because of the co- given in microns (urn); ratios (i.e., SW/TL) are occurrence of the new species with E. pennaki expressed in percent of the total length (TL) meas- Higgins, 1960, the description of which reflects ured on the midline, from the anterior margin of my own inexperience at the time, I shall redescribe segment 3 (first trunk segment) to the posterior it as well. The description of E. pacificus Schmidt, margin of segment 13, exclusive of spines. Maxi- 1974 is included because it closely resembles the mum sternal width (MSW) is measured at the new species from San Juan Island. Finally, while anteroventral margin of the widest pair of sternal studying the taxonomic history of E. dujardinii, I plates as first encountered in measuring each seg- discovered that a name I proposed for a species ment from anterior to posterior. Sternal width at from the Red Sea was preoccupied; therefore, a segment 12 (SW), or standard width, is measured substitute name is offered for E. brevispinosus Hig- at the anteroventral margin of the 12th sternal gins, 1966a. plates. METHODS.—Specimens were preserved in 70 per- Middorsal spines (D), lateral spines (L), and cent ethyl alcohol, 5 percent formalin, or Duboscq lateral accessory appendages (LA) are numbered by and Brasils fluid, and then transferred to a 70 segment and their cumulative mean length ex- percent ethyl alcohol-5 percent glycerin solution pressed by Dm, Lm, and LAm, respectively. Meas- that was allowed to evaporate to glycerin. Most urements are given for the lateral terminal spines specimens were removed from the glycerin and in- (LTS), lateral terminal accessory spines (LTAS), dividually placed in Hoyer's mounting medium, midterminal spine (MTS), and penis spines (P) in between two coverslips, and positioned on Cobb males. The first penis spine (P-l) is usually the aluminum slide frames. This mounting procedure anteriormost of three such spines and is dorsally allows the slide to be placed on either of its sur- displaced; the second (P-2) is usually the shortest faces so that both dorsal and ventral aspects of the and often more truncate (probably the functional specimen can be observed. penis); the third (P-3) is usually adjacent to the Hoyer's medium is necessary to soften the speci- second or slightly posterior to it. Both P-2 and P-3 men so that, by judicious manipulation of the are best observed in ventral aspect. coverslip, the specimen will assume a dorsoventral Several lateral spines appear to function as ad- position; this medium also clears the specimen, thus hesive tubes. This study will introduce the adhesive revealing the detailed structure of the exoskeleton. tubes of the fourth segment (L-4) as homologues A disadvantage of the Hoyer's medium is its of other lateral spines. Appendages that function tendency to clear the specimen too much, espe- as adhesive tubes will be noted in the appropriate cially over a period of several years. This may be section of the text. NUMBER 248
Specimens mentioned in this paper are deposited Stockholm, and the late Dr. B. G. Chitwood were in the National Museum of Natural History, Smith- helpful in this study. Dr. Peter Schmidt, Institute sonian Institution, under the catalog numbers of fur Zoologie der RWTH, Aachen, generously pro- the old United States National Museum (USNM). vided type specimens of E. pacificus for my use. ACKNOWLEDGMENTS.—I am grateful to Mr. John Thanks also go to Prof. Peter Dohrn, Stazione C. Boykin, University of Washington, the first to Zoologica, Naples and to Dr. Jose Stirn, Marine collect the new species from San Juan Island and Laboratory, Portoroz, Yugoslavia, for making their recognize its taxonomic problems; and to our mu- facilities available to me. Drs. Raymond B. Manning tual friend and colleague from this same institu- and Horton H. Hobbs, Jr., were generous in their tion, Dr. Eugene N. Kozloff, who accepted the advice and consultation during the preparation of challenge of culturing the new species, and pro- this manuscript and, along with Dr. Richard S. vided me with specimens both of the new species Houbrick, in the reading of the final draft. I also and of E. dujardinii that have made this study wish to express my appreciation to Mrs. Eleanor possible. I also acknowledge the cooperation of Dr. Goldsmith, Mrs. Martha Brewster, and Mrs. Marie Jean Merriman, California State College at Sonoma, Wallace for their help in the preparation of this whose recent doctoral dissertation and publication manuscript. Finally, I am pleased to acknowledge have contributed to the knowledge of the new the Sumner Gerard Foundation and the Smith- species from San Juan Island. The personal com- sonian Foreign Currency Program for their finan- munications of Dr. Tor G. Karling, University of cial support.
Key to Adults of the Genus Echinoderes 1. Middorsal spines absent 2 Middorsal spines present 5 2. Lateral spines (except for terminal segment) absent Echinoderes maxwelli (Omar-Cooper, 1957) Lateral spines present 3 3. Lateral spines on segments 7, 10, and 12 only Echinoderes bengalensis (Timm, 1958) Lateral spines absent on segment 12 4 4. Lateral spines on segments 7, 10, and 11 only Echinoderes caribiensis Kirsteuer, 1964 Lateral spines on segments 7 and 10 only, segments 3-4 expanded laterally Echinoderes capitata (Zelinka, 1928) 5. Middorsal spines on segments 6-10 only 6 Middorsal spines with other arrangement IS 6. Lateral spines on segments 7-12 only 7 Lateral spines with other arrangement 21 7. Lateral accessory spine on segment 10 Echinoderes dujardinii Claparede, 1863 Lateral accessory spines absent on segments 3-11 8 8. Trunk length greater than 300 /un 9 Trunk length less than 300 Mm 11 9. Lateral spine on segment 12 short (12-17 Mm), less than half the length of lateral spine on segment 11 Echinoderes pacificus Schmidt, 1974 Lateral spines on segment 12 nearly equal to length of lateral spines on segment 11 10 10. Lateral terminal accessory spines 12%—15% of trunk length, midventral placid wider than adjacent placids Echinoderes koxloffi, new species Lateral terminal accessory spines about 8% of trunk length, midventral placid not wider than adjacent placids Echinoderes pilosus Lang, 1949 11. Middorsal spine on segment 10 slightly longer than middorsal spine on segment 9 Echinoderes ehlersi Zelinka, 1913 Middorsal spine on segment 10 twice as long as middorsal spine on segment 9 12 12. Lateral terminal accessory spines about 41% of trunk length, no lateral accessory spine on segment 12 Echinoderes worthing Zelinka, 1928 Lateral terminal accessory spines about 23% of trunk length, lateral accessory spine ad- jacent to lateral spine 12 Echinoderes ferruginous Zelinka, 1928 13. Middorsal spine on segment 11 (juvenile?) Echinoderes sonadiae Timm, 1958 Middorsal spine absent on segment 11 14 4 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
14. Middorsal spine on segment 7 15 Middorsal spine absent on segment 7 18 15. Middorsal spines on segments 6, 7, and 9 only Echinoderes setigera Greeff, 1869 Middorsal spines with other arrangement 16 16. Middorsal spines on segments 6-9 only Echinoderes druxi d'Hondt, 1973 Middorsal spine on segment 5 17 17. Middorsal spines on segments 5-8 only Echinoderes tchefouensis Lou, 1934 Middorsal spines on segments 5-10 only Echinoderes borealis Greeff, 1869 18. Middorsal spines on segments 6 and 9 only Echinoderes citrinus Zelinka, 1928 Middorsal spines on segments 6, 8, and 10 only 19 19. Lateral accessory spines on segments 3, 8-10 with subdorsal spine on segment 3, terminal tergal extensions evenly tapering to point Echinoderes newcaledoniensis Higgins, 1967 Lateral accessory spine on segment 10 or missing on all segments except terminal segment of female, terminal tergal extensions with uneven margins 20 20. Lateral accessory spine on segment 10 only, middorsal spines short, (15-18 /*m). Terminal tergal extensions mesially recurved and somewhat rectangular with small protuberances at corners Echinoderes riedli Higgins, 1966a Lateral accessory spines absent on all segments except terminal segment of female, mid- dorsal spines long (73-125 ^m). Terminal tergal extensions rounded with elongate central spine Echinoderes arlis Higgins, 1966b 21. Posterior border of 12th tergal plate interrupted medially to form broad spine-shaped process. Small lateral spine or prominent hair (about 8 ^m) on segment 12, similar spine or hair on segment 13 anterior to lateral terminal accessory spine of female Echinoderes bookhouti Higgins, 1964b Posterior border of 12th tergal plate even. Lateral spine or prominent hair absent on segment 12, segment 13 with lateral terminal accessory spine of female and lateral terminal spine only 22 22. Lateral spine on segments 7-11 only 23 Lateral spines with other arrangement 25 23. Lateral terminal spines short, about 17% of trunk length. Spinous fringe most prominent at posteroventral margin of segment 4 and at lateral margins of sternal-tergal junctions of segments 9 and 10 Echinoderes brevicaudatus, substitute name for Echinoderes brevispinosus Higgins, 1966a Lateral terminal spines 24%-40% of trunk length, spinous fringe uniform on each segment where it occurs 24 24. Terminal sternal plates pointed, tergal plate extensions displaced medially from lateral terminal spines, midventral placid expanded at anterior margin, prominent spinous fringe along entire posteroventral margin of segment 3 ... Echinoderes remanei (Blake, 1930) Terminal sternal plates rounded, tergal plate extensions displaced laterally alongside lateral terminal spines, midventral placid evenly rounded or slightly truncate at anterior margin, spinous fringe more prominent near midline of posteroventral margin of segment 3 Echinoderes pennaki Higgins, 1960 25. Lateral spines on segments 6-9 only Echinoderes canariensis Greeff, 1869 Lateral spines with other arrangement 26 26. Lateral spines on all segments (probably error in description) Echinoderes steineri (Chitwood, 1951) Lateral spines on segments 8-11 only 27 27. Lateral terminal spines about 50% of trunk length, lateral spine on segment 11 same length as or slightly longer than that of segment 10 Echinoderes elongata (Nyholm, 1947b) Lateral terminal spines equal to or longer than trunk length, lateral spine of segment 11 very long Echinoderes levanderi Karling, 1954
Echinoderes dujardinii C llaparede "Echinoceras" Leuckart, 1854:355 [erroneous spelling; Helgoland]. FIGURES 1-12 Echinoderes Dujardinii Claparede, 1863:90-92, pi. 16: figs. 7-13 [type-locality: St. Vaast la Hougue].—Leuckart, 1869: "I'Echinodere" Duprdin, 1851:158, pi. 3: figs. 1-4 [St. Malo].— 300.—Greeff, 1869:88-89, pi. 4: figs. 1-5 [by inference: Schultze, 1853:253. Ostende, Xieuwpoort, Dieppe, Canary Islands; incorrectly NUMBER 248
cites Dujardin as author].—Panceri, 1876:4, 5, figs. 6, 7 Habroderes meridionales Zelinka, 1928:254, pi. 2: figs. 11-13 [Ischia, incorrectly cites Greef (sic) as author].—Reinhard, [synonomy].—Remane, 1936:333 [noted as larval stage of 1881:589 [Odessa, incorrectly cites Dujardin as author].— Echinoderes dujardinii]. Moebius, 1884:7, 1887:117.—Reinhard, 1885:5.—Cams, Centropsis parallela Zelinka, 1928:269, pi. 1: figs. 4, 5, 11, 12 1885:184.—Ludwig, 1886:882.—Reinhard, 1887:438 [incor- [Naples, Trieste].—Remane, 1936:332, 333, fig. 273 [noted rectly cites Dujardin as author].—Perrier, 1893:453.— as larval stage of Echinoderes dujardinii]. Zelinka, 1896:199 [incorrectly cites Greeff as author].— Echinoderes Masudai Abe, 1930:42, 43, pi. 1: figs. 1, 2 [Hiro- Grobben, 1905:336.—Schepotieff, 1907b:297, 298, pi. 17: shima.].—Remane, 1936:345, fig. 277.—Tokioka, 1949:67. figs. 1-6, 16a, pi. 18: figs. 22, 23, pi. 20: fig. 13 [Bergen, Echinoderes sieboldii.—Lou, 1934:3 [erroneous spelling]. Naples, Brindisi, Rovignj].—Southern, 1914:69, 71 [Clew Echinoderes masudai.—Lang, 1949:17.—Higgins, 1960:89 Bay, Blacksod Bay].—Zelinka, 1928:221 [lapsus calami}].— [nomen dubium]; 1966a:123; 1967:75; 1971:26. Abe, 1930:39-43. Echinoderes aff. dujardini.—Bacescu et al., 1963:137, 138, figs. Echinodera Dujardinii Gosse, 1864:403-404, pi. 2: fig. 16 [de- la-c [Agigea, probably not Echinoderes dujardinii]. cription taken from Dujardin, 1851 only].—Zelinka, 1928: Not Echinoderes brevispinosus Higgins, 1966a:l 18-121, figs. 1, 228 [synonomy]. 2; 1966b:519.—Schmidt, 1974:14. [=Echinoderes brevicau- Not Echinoderes Dujardinii.—Metschnikoff, 1865:459-461 datus, substitute name]. [Helgoland, =Echinoderes subfuscus Zelinka, 1928]; 1869: Not Echinoderes dujardinii.—Chitwood, 1964a:2 [Tomales 190-193 [Selerno, = Echinoderes subfuscus Zelinka, 1928]. Bay, Calif; identity uncertain]. Echinoderes brevispinosa Metschnikoff, 1869:190 [Salerno].— Not Echinoderes dujardini.—Merriman and Corwin, 1973: Leuckart, 1869:300-301.—Reinhard, 1887:438. 227-243, figs. 1-13 [San Juan Island, Washington; =Echino- Echinoderes Sieboldii Pagenstecher, 1875:117-123, pi. 7: figs. deres kozloffi, new species]. 1-6 [Palma]; 1877:88-89, fig. 69.—Cams, 1885:185.— Reinhard, 1887:438 [Odessa].—Schepotieff, 1907b:298, 299, REDESCRIPTION.—Adults (Figures 1-6), trunk pi. 17: fig. 7.—Abe, 1930:39-42. length, 328-405 um; MSW-9, 78-85 urn, 19.9-24.9 Echinoderes' brevispinosus.—Panceri, 1876:4 [correction of percent of trunk length; SW, 70-83um, 18.9-24.9 E. brevispinosa Metschnikoff, 1869].—Zelinka, 1928:228 [synonomy]. percent of trunk length. Echinoderes meridionales Panceri, 1876:5, 6, fig. 8 [Ischia].— Second segment with 16 anteriorly rounded Carus, 1885:185.—Zelinka, 1928:254, pi. 2: figs. 11-13 placids, midventral placid truncate, expanded [synonomy, =Habroderes meridionales']. slightly at anterior margin, distinctly larger than Echinoderes dujardiniiQ).—Hartog, 1896:236, fig. 120 [Wor- adjacent placids; trichoscalid plates on sides of mid- thing].—Mclntyre, 1962:503. ventral placid with medial indentation on anterior Echinoderes dujardini.—Hartog, 1896:237.—Remane, 1928:75, 76, figs. 13, 15 [incorrectly cites Greeff as author on p. 76 margin, posterior margin expanded laterally. and in figure legends].—Johnston, 1938:5.—Higgins, 1960: Segments 3-12 with short hairs, pattern distinc- 88-90.—Mclntyre, 1962:503.—Higgins, 1964a: 244, 246; tive (Figures 1, 2); posterior border of segments 1964b:489-491; 1966a:120; 1967:75,79; 1969b:113; 1971:25.— 5-11 with fine pectinate fringe ventrally, segments Kozloff, 1972:121.—Moore, 1973:349.—Higgins, 1974:512.— 6-11 with pectinate fringe dorsally; terminal seg- Boykin, 1974:40.—Schmidt, 1974:12. Echinoderes dujardinii.—Levander, 1900:20.—Schepotieff, ment with few hairlike processes along posterior 1907a: 147, 148, figs. 10-12 [Bergen, Naples].—Zelinka, 1907: margin, posterior margin of tergal plate deeply 132; 1908:630-641, figs. 1, 5, 6, 9 [incorrectly cites Greeff as incised forming pointed extensions mesial to base author]; 1912:520-527 [incorrectly cites Greeff as author]; of each lateral terminal spine, sternal plates broadly 1913:419, 420 [incorrectly cites Greeff as author]; 1928:221 rounded with spinous extensions, 10-12 urn in [incorrectly cites Greeff as author], 228 [cites Greeff as author in synonomy], 228-235, figs. 17-19; pi. 3: figs. 1, length (Figures 3, 5). 2, pi. 10: figs. 1-5, 7-23 [Naples, Trieste, Kiel].—Remane, Middorsal spines on segments 6-10, increasing 1928:60, 62, figs. 7, 9 [incorrectly cites Greeff as author].— uniformly in length, 13-29 um; lateral spines on Lou, 1934:3.—Remane, 1936: 333, figs. 215, 252, 266.— segments 4 and 7-12, 14-29 um in length; L-4, 7, Zaneveld, 1938:261 [Scheveningen].—Nyholm, 1947a:424 10 and 12 each with adhesive gland at base; L-10 [Gullmar Fjord]; 1947b:5, fig. 1.—Lang, 1949:3, 17.— Tokioka, 1949:67.—Hyman, 1951:180.—Karling, 1954:189- accompanied by dorsally adjacent lateral accessory 192.—Kirsteuer, 1964:389—Marinov, 1964:62, 63, fig. 2 spine of nearly equal length, 21-26 ^m; L-4 usually [Varna, doubtful record]. longer (mean 24.2 um) than remaining lateral Echinoderes sieboldi.—Zelinka, 1928:228 [synonomy].— spines; lateral terminal spines long, 160-192 um, Remane, 1936:344 [synonomy]. 40.7-54.8 percent of trunk length; lateral terminal Habroderes dujardinii Zelinka, 1928:248-250, pi. 4: figs. 1, 2 [described as "n. 1. (zu Echinoderes dujardinii Clap.)"].— accessory spines of female 52-62 um in length; males Remane, 1936:332, 333, fig. 273 [noted as larval stage of without lateral terminal accessory spines but with Echinoderes dujardinii]. penis spines in same position, anteriormost penis SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
100pm
KI
1 2 FIGURES 1-2.—Echinoderes dujardinii, adult female (USNM 53342, RH125.14), neck and trunk segments: 1, ventral view; 2, dorsal view. NUMBER 248
50pm
6 FIGURES 3-6.—Echinoderes dujardinii, segments 12-13: 3, ventral view, lateral half, adult female (USNM 53342, RH125.14); 4, dorsal view, lateral half of same adult female; 5, ventral view, lateral half, adult male (USNM 53342, RH125.35); 6, dorsal view, lateral half of same adult male. spine (P-l) 23-33 um in length, mesially adjacent Second segment similar to that of adult, both penis spine (P-2) 24-36 um, posteriorly adjacent placids and trichoscalid plates less well developed. penis spine (P-3) 26-39 um in length. Trunk segments with fewer hairs than adult, Pachycycli (thickened anterior margins of trunk pattern less distinctive and more variable; posterior segments) well developed, forming a distinctive pat- borders of segments without pectinate fringe, with tern at ventral midline and at attachments of lateral hairs (striations?) along border, group of prominent terminal spine muscles on segment 13; distinctive hairs (striations?) at ventral midline, terminal seg- muscle scars on ventrolateral portion of first trunk ment slightly incised dorsally and ventrally, sternal segment, similar (but reversed orientation) lateral area without spinous extensions, tergal area with scars on eighth sternal plates; sensory spots, 2-3 um small extensions 2-4 um in length, not evenly in diameter, situated middorsally on segments 3-5, tapered. with two such spots on segment 12 (possibly one on Middorsal spines on segments 6-11, increasing each of segments 12 and 13), dorsolaterally on seg- uniformly in length, 17-39 um; lateral spines on ments 6-11, and ventromesially on segments 5-12 segments 4 and 7-12, 17-26 jun in length; L-4, 7, (Figures 1, 2). 10 and 12 with poorly developed adhesive glands Morphometric data for adult specimens are at base; L-10 and L-12 accompanied by a dorsally shown in Table 1. adjacent lateral accessory spine of nearly equal JUVENILE STAGES.—Preadult stage (J-6, "Habro- length, 21-25 um, but thinner; L-4 more prominent deres-stage," Figures 7, 8) trunk length, 320-328 um; than remaining lateral spines; lateral terminal estimated MSW-9, 78-80 (im, 24.3-25.6 percent of spines long, 148-152 um, 45.1-48.7 percent of trunk trunk length; estimated SW, 65-72 um, 19.8-23.5 length; lateral terminal accessory spines 42-47 ^m percent of trunk length (estimated since tergal- in length, sexes often indistinguishable in juvenile sternal junctions are not defined in juvenile stages). stages unless developing oocytes visible. SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE 1.—Measurements (/im) and indices (%) for Echinoderes dujardinii adults
Standard Standard Coefficient Standard Standard Coefficient
TL a ... 26 328-380 350.5 15.4 3.0 4.3 L-4 (AT) 16 17-26 23.0 2.3 0.6 9.8 20 336-405 367.9 21.9 4.9 6.0 9 ... 17 22-28 25.2 1.6 0.4 6.2 at... 46 328-405 358.0 18.3 2.7 5.1 cT9 ... 33 17-28 24.2 1.9 0.3 7.9
SU SW/TL HSW-9 MSH/TL a 26 20.7-24.9 22.7 1.3 0.2 5.5 L-10 Om Dm/TL D-6 a ... 11 13-16 14.0 1.0 0.3 7.1 L-12 a .. 26 20-29 23.0 1.9 0.4 8.2 9 ... 19 13-16 14.2 1.1 0.3 8.0 9 .. 19 20-28 24.4 2.1 0.5 8.5 tf9 ... 30 13-16 14.1 1.1 0.2 7.6 cf9 .. 45 20-29 23.6 1.9 0.3 8.2 0-7 D-8 cf ... 18 11-23 16.3 1.1 0.3 7.0 LTS/TL cf 26 44.3-54.8 50.2 2.9 0.6 5.7 9 ... 19 14-21 17.1 1.7 0.4 9.7 9 .. 20 40.7-54.0 47.9 3.8 0.8 7.8 16.7 1.4 0.2 8.9 cf9 ... 37 11-23 <*9 .. 46 40.7-54.8 49.2 3.0 0.5 6.6 D-9 cf ... 20 13-23 18.6 2.6 0.6 13.9 LTAS 9 .. T9 52-62 56.6 3.1 0.7 5.4 9 ... 19 16-23 19.5 2.7 0.6 14.0 <*9 ... 39 13-23 19.0 2.6 0.4 13.7 LTAS/TL 9 .. 19 13.7-17.5 15.3 1.2 0.3 7.8 D-10 tf ... 23 20-26 22.6 2.4 0.5 10.4 P-l a .. 25 23-33 27.4 2.9 0.6 10.5 9 ... 20 16-27 21.9 3.1 0.7 14.0 <*9 ... 43 16-27 22.3 2.7 0.4 12.0 P-2 Lm 18 17.7-22.8 20.6 1.4 0.3 6.9 P-3 a 19 26-39 33.5 3.9 0.9 11.5 9 ... 17 19.2-25.0 21.2 1.5 0.4 6.8 •*•«... 35 17.7-25.0 20.9 1.4 0.2 6.8 Lm/TL C ... 18 4.6-6.7 5.8 0.5 0.1 8.5 9 ... 17 4.9-6.9 5.8 0.1 0.6 9.9 Pachycydi not well defined, muscle scars on ven- preadult, more scattered; terminal segment slightly trolateral areas of first trunk segment reduced, no incised ventrally with suggestions of pointed tergal scars on segment 8; sensory spots, 2-3 um in diam- extensions. eter, situated middorsally on segments 4-5 and Middorsal spines on segments 6-11, D-6-10, 12-13, laterodorsally on segments 6-13, mesially on 14-39 \tsn, increasing uniformly in length to seg- sternal plates 4-11. ment 11, D-ll, 79 um, twice the length of D-10, Fourth-stage juvenile (J-4, "Habroderes-stage," extending slightly beyond terminal segment; lateral Figures 9, 10) trunk length, 224 jun; estimated spines similar to preadult stage, 16-26 um in length; MSW-9, 72 urn, 32 percent of trunk length; esti- lateral terminal spines 160 um in length, 71.4 per- mated SW, 65 um, 29 percent of trunk length. cent of trunk length; lateral terminal accessory Second segment as in preadult, less well devel- spines 47 um. oped. Third-stage juvenile (J-3, "Hapaloderes-stage," Trunk segments similar to those of preadult but Figures 11, 12) trunk length, 200-208 um; estimated not well defined posteriorly so that only 10 trunk MSW-9, 72 um, 34.3 percent of trunk length; esti- segments apparent; series of medial hairs (stria- mated SW, 51 (wn, 24.5 percent of trunk length. tions?) both dorsally and ventrally indicating area Second segment as in J-4 stage, less well devel- of presumptive segment 12; corresponding area of oped. segment 13 less distinct; hairs less obvious than in Trunk segments similar to those of J-4 stage, less NUMBER 248 ,V (A. 100pm , -J _ fj}ii MA \ . ' 1 FIGURES 7-8.—Echinoderes dujardinii, preadult (J-6) stage (USNM 53345, RH125.49), neck and trunk segments: 7, ventral view; 8, dorsal view. 10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY 100pm FIGURES 9-10.—Echinoderes dujardinii, juvenile J-4) stage (USNM 53344, RH125.53), neck and trunk segments: 9, ventral view; 10, dorsal view. NUMBER 248 11 100pm I , f I I FIGURES 11-12.—Echinoderes dujardinii, juvenile (J-3) stage (USNM 53343, RH125.54), neck and trunk segments: 11, ventral view; 12, dorsal view. well defined posteriorly, striations along posterior tions incomplete on presumptive segment 10, re- margins of segments 3-9 defining segments, stria- stricted to midventral area, striations restricted to 12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY middorsal and midventral areas of presumptive seg- to the trunk length, 77 percent as contrasted with ment 11, striations apparent on midventral area of 41-55 percent in E. dujardinii. Echinoderes fer- presumptive segment 12; terminal segment blunt, rugineus Zelinka, 1928 has been found with E. du- without extensions. jardinii at both Naples and Trieste (Zelinka, 1928). Middorsal spines on segments 6-12 (D-12 essen- Echinoderes ferrugineus also is smaller than E. du- tially midterminal), D-6-9, 17-29 um, increasing jardinii and has longer lateral terminal spines uniformly in length to segment 10, D-10, 39-43 relative to its trunk length, about 67-77 percent. urn in length, D-ll, 98-109 urn in length, D-12 Echinoderes ferrugineus differs from E. ehlersi (midterminal) 136-170 um in length, 60.7-85.0 per- in that D-10 in the former species is 55 urn, nearly cent of trunk length; lateral spines similar to J-4 twice the length of D-9; in E. ehlersi, the mid- stage except LA-12 absent, 18-22 um in length; dorsal spines increase uniformly from 8-14 um in lateral terminal spines 60-64 \im, 28.8-30.0 percent length. of trunk length; lateral terminal accessory spines Echinoderes worthingi Zelinka, 1928 also occurs 46 um. with E. dujardinii. One specimen was collected along with E. dujardinii in this study and Southern MATERIAL EXAMINED.—46 adults (USNM 53342) consisting of 26 males and 20 females, 4 preadults (J-6, "Hapaloderes- (1914) found both species at Blacksod Bay, Ireland. stage") (USNM 53345), one fourth-stage juvenile (J-4, "Hapa- Echinoderes worthingi is similar in size and other loderes-stage") (USNM 53344), and 2 third-stage juveniles characteristics to both E. ehlersi and E. ferrugineus (J-3, "Habroderes-suge") (USNM 53343); col. Dr. Eugene N. as noted by Zelinka (1928). Echinoderes worthingi Kozloff, 19 October 1973, Roscoff, Nord-Finistere, France most closely resembles E. ferrugineus in that the (from the surface of sandy mud in the port). length of D-10, 45-50 um, is twice that of D-9, REMARKS.—Dujardin (1851) and many subse- 19-23 um. The two species appear to be distinguish- quent authors prior to Zelinka (1928) introduced able by the shape of the terminal segment, a slightly certain ambiguities into the description of Echino- smaller lateral terminal spine, and the presence of deres dujardinii. The most common problems in- a prominent sensory hair (spine?) posteriorly ad- clude the interpretation of the animal's length and jacent to the lateral spine on segment 12 in E. fer- the position of the spines on the trunk segments. rugineus; a similar hair exists in E. pacificus Dujardin, for example, probably included the head (Figures 28, 30). and neck and may have included some juvenile Three additional species with the D-6-10, L-A, stages in his total length measurements (300-550 7-12 spine combination are also similar to E. um). Prior to Zelinka (1928), most authors were not dujardinii in size. Echinoderes pilosus (from South consistent in numbering segments; consequently, Georgia Island) is poorly described, and as men- one must carefully compare the illustrations with tioned previously, I am including it because there the text when assessing the correct position of mid- is a chance that Lang (1949) overlooked the lateral dorsal and lateral spines. spines (= adhesive tubes) on segment 4. Echino- The most accurate illustration of E. dujardinii deres pilosus differs from E. pacificus Schmidt, published prior to Zelinka (1928) was that of Greeff 1974 (from the Galapagos) in that the former has (1869). One of the most important characteristics of relatively longer lateral terminal spines, about 52 this species, the presence of two lateral spines on percent of the trunk length compared with 27-36 segment 10, was noted by Dujardin (1851), but percent in the latter. If Lang is correct, a distin- ignored by many subsequent authors. guishing feature of E. pilosus is the uniform size of Including E. dujardinii, the adults of seven spe- the placids. In the other species, however, the mid- cies of Echinoderes have middorsal spines on seg- ventral placid is much wider than the adjacent ments 6-10 and lateral spines on segments 4, 7-12 ones. Both E. pacificus and the remaining species, (£. pilosus Lang, 1949 may have lateral spines or E. kozloffi, new species, are discussed more exten- adhesive tubes on segment 4 although they are not sively below. included in the author's description). Echinoderes All six species mentioned in the foregoing dis- ehlersi Zelinka, 1913 (from Zanzibar) is 228 um cussion differ from E. dujardinii by their lack of a long, much smaller than E. dujardinii, and has sig- lateral accessory spine on segment 10. nificantly longer lateral terminal spines in relation Two additional species of Echinoderes share with NUMBER 248 13 E. dujardinii the presence of a lateral accessory Echinoderes kozloffi, new species spine on segment 10. These include E. riedli Hig- gins, 1966a (from the Red Sea) and E. newcale- FIGURES 14-21 doniensis Higgins, 1967. Echinoderes riedli is Echinoderes.—Kozloff, 1972:121, figs. 1-18—Higgins, 1974:514, smaller, 238 \un in length. Echinoderes newcale- figs. 11-16. doniensis is the only member of this genus having Echinoderes dujardini.—Merriman and Corwin, 1973:227-243, lateral accessory spines on segments 8-11 and is figs. 1-13. unique in possessing dorsolateral spines on segment Echinoderes sp.—Boykin, 1974:40. 4. Both E. riedli and E. newcaledoniensis differ DIAGNOSIS.—Trunk length, 328-376 jim, mid- from E. dujardinii by having middorsal spines on dorsal spines on segments 6-10, increasing uni- segments 6, 8, and 10 only. formly in length; lateral spines on segments 4, 7-12, DISTRIBUTION.—Echinoderes dujardinii has been with adhesive glands at base of L-4, 7, 10 and 12, reported from 26 localities (Figure 13, also see an- lateral spine of segment 12, 20-30 um in length; notations in synonomy). Most of the reports are lateral extensions of terminal tergal plate tapering European and include both the northern and evenly to point protruding about 5 um beyond southern coastal areas of the continent. These dis- extensions of tergal plate (exclusive of prominent tribution records are probably reliable except for spines on latter). those from the Black Sea. Based on the evidence DESCRIPTION.—Adults (Figures 14-19), trunk presented by Reinhard (1881), Bacescu et al. (1963), length, 328-376 um; MSW-9, 68-74 um, 18.0-22.3 and Marinov (1964), I believe these records are percent of trunk length; SW, 62-69 urn, 17.5-20.0 questionable. Similarly, the reports of E. dujardinii percent of trunk length. from Japan (Tokioka, 1949) and the Pacific coast Second segment with 16 anteriorly rounded plac- of the United States (Chitwood, 1964a; Merriman ids; midventral placid truncate, not expanded lat- and Corwin, 1972) are based on misidentifications erally at anterior margin, distinctly larger than of E. kozloffi, new species. adjacent placids; trichoscalid plates on sides of FIGURE 13.—Echinoderes dujardinii, distribution records, open circles indicate doubtful records. 14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY '/r^/ 100pm ^iffi ""»«*,. te'ito wu 14 15 FIGURES 14-15.—Echinoderes kozloffi, new species, holotypic female (USNM 5SS37), neck and trunk segments: 14, ventral view; 15, dorsal view. NUMBER 248 15 midventral placid indented at anteromedial margin, terminal accessory spines of female 46-59 um in posterior margin expanded laterally, four tricho- length; males without lateral terminal accessory scalid plates on dorsal placids much smaller than spines but with penis spines in same position; those on ventral placids. anteriormost penis spine (P-l) 25-33 um in length, Segments 3-12 with short hairs, pattern distinc- mesially adjacent penis spine (P-2) 21-27 um in tive (Figures 14, 15); posterior border of segments length and distinctly broader, posteriorly adjacent 4-11 with fine pectinate fringe dorsally and ven- penis spine (P-3) 35-42 um in length, generally trally; terminal segment with few hairlike processes tending to curve mesially, crossing lateral terminal along posterior margin; posterior margin of tergal spine. plate deeply incised, laterally forming pointed, Pachycycli well developed, forming distinctive evenly tapered extensions mesial to base of each pattern at ventral midline and superficial to lateral lateral terminal spine; sternal plates broadly terminal spine muscle attachments on segment 13; rounded with spinous extensions, 15-22 um in length (Figures 16, 18). muscle scars on ventrolateral portion of first trunk Middorsal spines on segments 6-10, increasing segment almost indistinguishable, similar V-shaped uniformly in length posteriorly, 20-45 um; lateral scars on either side of ventral midline of segments spines on segments 4 and 7-12, 24-32 um in length; 4-9; prominent scars on lateral margins of sternal L-4, 7, 10 and 12 with adhesive gland at base; L-8 plates 8-9, more centrally located on sternal plates usually shorter (mean 22.9 um) than L-7 (mean 27.5 of segment 10 (Figure 14); sensory spots, 2-3 um in um) or L-9 (mean 26.1 um); L-9-12 all similar in diameter, situated middorsally on segments 3—5 length, 22-32 um; lateral terminal spines long, 144- with two such spots on segment 12 (possibly one on 180 um, 42.0-52.4 percent of trunk length; lateral each of segments 12 and 13), dorsolaterally on seg- 50pm 18 19 FIGURES 16-19.—Echinoderes kozloffi, new species, segments 12-13: 16, ventral view, lateral half, holotypic female (USNM 53337); 17, dorsal view, lateral half, holotypic female; 18, ventral view, lateral half, allotypic male (USNM 53338); 19, dorsal view, lateral half, allotypic male. 16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY ments 3-11, ventromesially on segments 4-12, an- lateral to midline; terminal segment slightly incised terolaterally on ventral surface of segment 3, and dorsally and ventrally, sternal area without spinous possibly near lateral margin of sternal plates 5-7 extensions, tergal area with evenly tapered exten- (Figures 14, 15). sions, 2-4 um in length. Morphometric data for adult specimens are Middorsal spines on segments 6-11, increasing shown in Table 2. uniformly in length, 17-62 um (mean 27.9-34.0 urn, JUVENILE STAGES.—Preadult stage (J-6, "Hab- 10.7-12.0 percent of trunk length), D-ll extending roderes-stage," Figures 20, 21) trunk length, 276- beyond terminal tergal borders; lateral spines on 300 um; estimated MSW-9, 64-68 um, 23-24 per- segments 4, 7-12, 17-26 um in length; L-4, 7, 10, cent of trunk length; estimated SW, 62-64 um, and 12 with poorly developed adhesive glands at 22-23 percent of trunk length (estimated since base; L-12 accompanied by a lateral accessory spine tergal-sternal junctions not defined in juvenile situated dorsally to it, slightly longer (22-26 um) stages). and thinner; L-4 also thinner than other lateral Second segment similar to that of adult, both spines; lateral terminal spines long, 152-160 um, placids and trichoscalid plates less well developed. 50.7-56.5 percent of trunk length; lateral terminal Trunk segments with fewer hairs than adult al- accessory spines 35-40 um in length. though hairs as long as in adult (5-9 um), pattern Pachycycli not well defined, muscle scars similar less distinctive and more variable; posterior borders to those of preadult stage of E. dujardinii. of segments without pectinate fringe, with hairs Fifth-stage juvenile (J-5, "Habroderes-stage") (striations?) along border, group of prominent hairs trunk length, 224-256 um; other measurements sim- (striations?) at ventral midline and second group ilar to preadult stage described above. TABLE 2.—Measurements Qua) and indices (%) for Echinoderes kozloffi, new species, adults Standard Standard Coefficient Standard Standard Coefficient Char acter Number Range Mean Deviation Error of Variability Character Number Range Mean Deviation Error of Variability Tl a . 20 328-364 334.4 8.2 1.8 2.4 Lm/TL tt . 20 7.1-7.9 7.6 0 2 0.1 3.5 9 . 20 332-376 356.0 12.9 2.9 3.6 9 . 20 6.7-8.1 7.4 0 4 0.1 5.6 rf«. 40 328-376 350.2 12.2 1.9 3.5 O9. 40 6.7-8.1 7.5 0 3 0.1 4.5 SW a . 19 62-66 62.3 1.2 0.3 1.8 L-4 (AT)d- . 20 26-29 27.0 1 1 0.2 3.9 9 . 20 62-69 66.4 1.9 0.4 2.9 9 . 20 25-29 27.1 1 3 0.3 4.9 d9. 39 62-69 65.4 1.9 0.3 2.9 rf9. 40 25-29 27.0 1 2 0.2 4.4 SW/TL MSW-9 tt . 19 68-72 69.5 1.3 0.3 l.S 1-8 a ., 20 21-25 22.9 1 4 0.3 6.2 9 . 20 69-74 71.8 1.8 0.4 2.5 9 .. 20 21-26 22.8 1 3 0.3 5.8 ri. 39 68-74 70.7 1.9 0.3 2. tf9.. 40 21-26 22.9 1 3 0.2 5.8 MSW/TL tt . 19 18.8-21.4 19.9 0.8 0.2 3.J L-9 a .. 20 24-28 26.2 0 9 0.2 3.4 9 . 20 18.0-22.3 20.1 1.0 0.2 5.1 9 .. 20 23-29 26.0 1 3 0.3 5.0 cf9. 39 18.0-22.3 20.1 0.8 0.1 4.; <&.. 40 23-29 26.1 1 1 0.2 4.2 Di tf . 9 26.6-29.8 28.7 1.1 0.4 L-10 Dn/TL a . 9 7.6-9.1 8.3 0.5 0.2 5 L-ll a .. 20 23-30 26.6 1 7 0.4 6.4 9 . 11 7.1-8.1 7.6 0.3 0.1 4. 9 .. 20 25-32 27.8 1 7 0.4 6.3 rfV. 20 7.1-9.1 7.9 0.5 0.1 6. D-6 a . 10 20-26 23.3 2.0 0.6 8.1 L-12 a .. 20 22-29 25.6 1 9 0.4 7.5 9 . 14 20-26 22.0 1.9 0.5 8. 9 .. 20 23-32 26.5 2 4 0.6 9.2 a 9. 24 20-26 22.5 2.0 0.4 8. 09.. 40 22-32 26.0 2 2 0.4 8.5 a . 13 21-27 24.4 1 .8 0.5 4.5 D-7 5' LTS a .. 20 144-172 157.8 7 1 1.6 12 22-26 23.4 1.2 0.3 4.0 9 . 6. 9 •• 20 160-180 170.1 6 7 1.5 d9. 25 21-27 23.9 1.6 0.3 O9-. 40 144-180 164.0 9 2 1.5 5.6 D-8 a . 13 23-:? 25.7 1.0 0.3 4.0 LTS/TL 0 .. 20 40.0-50.0 45.9 2 5 0.6 5.5 9 • 15 22-26 24.4 1.7 0.4 6.8 9 •• 20 43.5-52.4 47.9 2 5 0.6 5.2 tf9. 28 22-27 25.0 1.5 0.3 6.0 O9. 40 42.0-52.4 46.9 2 7 0.4 5.7 D-9 d" . 14 26-30 27.6 1.3 0.4 4.9 LTAS 9 .... 20 46-59 52.6 3.5 0.8 6.7 9 . 16 24-33 26.9 2.2 0.5 8.1 cf9. 30 24-33 27.2 1.8 0.3 6.7 LTAS/Tl V 20 12.6-15.8 14.8 1.0 0.2 6.7 D-ia cf . 20 38-45 41.6 2.5 0.6 6.1 P-l In rf . 20 24.7-27.7 26.0 0.8 0.2 2.9 P-3 a 18 35-42 38.3 1.9 0.4 4.9 9 . 20 24.0-27.7 26.4 0.9 0.2 3.5 O9. 40 24.0-27.7 26.2 0.9 0.1 3.3 NUMBER 248 17 100pm 20 21 FIGURES 20-21.—Echinoderes kozloffi, new species, preadult (J-6) stage (USNM 53341, RH145.43), neck and trunk segments: 20, ventral view; 21, dorsal view. 18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY Middorsal spines on segments 6-11, increasing sternal plates of the terminal segment distinguish uniformly in length, 15-72 um (mean 33.0-39.8 the two species. The mesial margins of the terminal um), 14.7-15.8 percent of trunk length. Note: This sternal extensions of E. pacificus are interrupted and latter statistic distinguishes J-5 from J-6. beset with a series of prominent hairs (Figure 28); HOLOTYPE.—Adult female, TL 344 urn; North in E. kozloffi these margins are even and have a less Bay, San Juan Island, Washington, USA (48°31.0'N, obvious series of hairs. Both species have hairs 123°O1.O/W); 9 August 1975; col. E. N. Kozloff; along the mesial margins of the terminal sternal USNM 53337. extensions but, again, those of E. pacificus are more ALLOTYPE.—Adult male, TL 348 um; as holotype; distinct, each hair associated with a steplike inter- USNM 53338. ruption of the margin. Also, the sternal extensions PARATYPES.—19 females and 19 males, TL 328- of both species have a spinous process. The proc- 376 um; data as for holotype; USNM 53339; 5 pre- esses in E. pacificus, however, are half the length adult juveniles, TL 224-300 um, 5 J-5? juveniles, (8-12 um) of those in E. kozloffi (17-25 um). Echi- 224-256 um; as holotype; USNM 53340; 4 females noderes pacificus is discussed later in this paper. and 4 males, TL 328-364 um; Reid Harbor, Stuart Echinoderes kozloffi is similar to E. pilosus. As / Island, Washington, USA (48°40.0 N, 123° 1 LOW); noted above, E. pilosus is not well described but 31 July 1963; col. J. C. Boykin; USNM 53347; 3 has the same general trunk length and spination preadult juveniles, TL 280-292 um; data as for formula as both E. kozloffi and E. pacificus. above; USNM 53348. Echinoderes pilosus appears to differ from E. koz- REMARKS.—Echinoderes kozloffi, new species, loffi in only two principal characters: E. pilosus most closely resembles E. pacificus from the Gala- has shorter lateral terminal accessory spines, about pagos Islands. I consider the two sibling species. 35 um, about 8 percent of the trunk length, and A similar sibling relationship involving the two the midventral placid is the same width as adjacent geographic areas has been noted for two species of placids. This latter character is unique within the otoplanid turbellarians: Philosyrtis sanjuanensis genus. Ax and Ax, 1967 (from San Juan Island, Washing- Echinoderes kozloffi is sympatric with E. pennaki ton) is a sibling species of P. santacruzensis Ax and Higgins, 1960 but differs in that the latter species Ax, 1974 (from the Galapagos Islands) according to lacks a lateral spine on segment 12, has conspic- these authors. Echinoderes kozloffi is similar to E. uously longer lateral spines (mean 58.1 um, 14 pacificus in total length, standard width, maximum percent of the trunk length), a prominent ventral sternal width, and general appearance. The range spinous fringe on the posterior border of the first of the mean lengths of middorsal spines in E. koz- trunk segment, lacks spinous processes on the loffi (25.4-29.8 um) and those of E. pacificus (46.5- terminal sternal plates, and has slightly longer, 56.5 yon) is one of the significant differences be- more pointed terminal tergal extensions. Other tween the two species. Other differences include the differences are mentioned in the discussion of longer (25-29 um) lateral spine on segment 4 in E. pennaki that follows. E. kozloffi, which contrasts with the shorter (12-17 ETYMOLOGY.—This species is named in honor of um) lateral spine on segment 4 in E. pacificus; in Eugene N. Kozloff, a fellow student of the the former species the lateral terminal spines, 144- Kinorhyncha. 180 um, 42.0-52.4 percent of the trunk length, are longer than those of the latter species, 90-118 um, 27.0-36.0 percent of the trunk length. The lateral Echinoderes pennaki Higgins terminal accessory spines of E. kozloffi are slightly longer (46-59 um) than those of E. pacificus (34- FIGURES 22-25 41 um). Echinoderes pennaki Higgins, 1960:86; 1961:81.—Chitwood, The relatively long (22-30 um) lateral spine on 1964b:3.—Higgins, 1964a:246; 1964b:479—Kirsteuer, 1964: segment 12 of E. kozloffi is a particularly notice- 389.—Higgins, 1966a: 120; 1966b:519; 1967:75.—Schmidt, able feature that contrasts with the short (12-17 1974:13.—Boykin, 1974:40. um), blunt lateral spine on segment 12 of E. pa- REDESCRIPTION.—Holotypic adult female (Figures cificus. The posterior margins of both tergal and 22-25), trunk length, 404 um; MSW-8, 62 um, 15 NUMBER 248 19 100pm FIGURES 22-23.—Echinoderes pennakt, holotypic female (USNM 29746), neck and trunk segments; 22, ventral view; 23, dorsal view. 20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY percent of trunk length; SW, 56 um, 14 percent of Middorsal spines on segments 6-10, increasing trunk length. uniformly in length, 48-70 um, mean length 59.6 Second segment with 16 anteriorly rounded um, 15 percent of trunk length; lateral spines on placids; midventral placid slightly truncate, dis- segments 7-11, L-4 (=adhesive tube) absent al- tinctly larger than adjacent placids; trichoscalid though a scar, or perhaps a sensory spot, located plates on sides of midventral placid indented slightly mesial to usual position of L-4; L-7 slightly on anterior margin, short and broad, shorter (14 um) than remaining lateral spines (32- trichoscalid plates on dorsal surface similar but 36 um) subequal in length, mean length of lateral equally as broad as long. spines 29.8 um, 7 percent of trunk length; lat- Segments 3-12 with prominent hairs (10-15 um), eral terminal spines long, 156 um, 39 percent of pattern distinctive (Figures 22-23); posterior border trunk length; lateral terminal accessory spines 52 of segments 3-11 with pectinate fringe both dor- um in length. sally and ventrally, fringe on ventral surface of Pachycycli well developed, forming distinctive segment 3 almost spinose, more finely pectinate on pattern at ventral midline and superficial to attach- remaining segments, slightly evident near posterior ment of lateral terminal spine muscles on segment border of 12th sternal plates; terminal segment 13; small muscle scars visible near midventral line with minute hairs on border of tergal and sternal of segments 3-4, 7, 11, and 12, more prominent scars plates, posterior margin of tergal plate deeply dorsolaterally on segments 3, 5-12; sensory spots sit- incised forming pointed extensions mesial to base of uated ventrolaterally on segment 3 and possibly on each lateral terminal spine, sternal plates broadly segment 4 (possibly adhesive tube openings?). rounded without spinous extensions (Figures MATERIAL EXAMINED.—4 adult females including holotype 24, 25). (USNM 29746) and 3 paratypes (USNM 29747); col. P. L. Illg, 16 July 1958, East Sound, Orcas Island, Washington, USA, from a depth of 32 meters. REMARKS.—Eighteen years of research on the Kinorhyncha has prompted me to reexamine my first species descriptions with better experience to guide me. The combination of this experience and better optical instruments has allowed me to illus- trate and describe E. pennaki more accurately. For example, the range of "total length" given in my original description (Higgins, 1960) is "390-430 um (taken along dorsal surface between anterior edge of second zonite and posterior edge of zonite 13)." 50pm More standard measurements are now taken from the anterior edge of segment 3 to the posterior edge of segment 13. Similarly, the maximum width of 80-90 um originally given for this species differs from the more precise and standardized measure- ment of maximum sternal width as noted in previ- ous publications (Higgins, 1967). Echinoderes pennaki most closely resembles E. remanei (Blake, 1930) redescribed by me (Higgins, 1964a). The two species differ in size: E. pennaki is larger (380-404 um) than E. remanei 282-358 um) yet both have the same spination formula (D-6-10, 24 L-4? 7-11). Both have a prominent spinous fringe FIGURES 24-25.—Echinoderes pennaki, segments 12-13 of holo- on the posteroventral border of segment 3; this typic female (USNM 29746): 24, ventral view, lateral half; 25, feature is more prominent in E. remanei and also dorsal view, lateral half. occurs on the fourth segment. The midventral NUMBER 248 21 placid of E. remanei is expanded laterally along its situated near the sensory spots of the 10th sternal anterior margin, and the border of the terminal plates. Sensory spots occur on either side of the sternal plates is pointed, not rounded as in E. pen- ventral midline of segments 4-12. Three spots are naki (Figure 24). Our understanding of both species present on the dorsal surface of the first trunk would benefit from an expanded study of their segment; only the median spot persists on the fol- taxonomic characters based on larger numbers of lowing two trunk segments (segments 4-5). Sensory individuals in a given sample. spots on the segments bearing middorsal spines are on both sides of the midline. Two middorsal sensory spots appear to be on segment 12; one of Echinoderes pacificus Schmidt these may be associated with segment 13. FIGURES 26-31 MATERIAL EXAMINED.—Holotypic male, TL 372 Mm (USNM Echinoderes pacificus Schmidt, 1974:1. 53335); 8 paratypic females and 7 paratypic males (USNM 53336); col. P. Schmidt, July-nSeptember 1972, Academy Bay, This species has been described with considerable Station IX, 5c, upper subtidal sediments. accuracy by my colleague, Dr. Peter Schmidt (1974). In studying the holotype and paratype material REMARKS.—As noted in the discussion of E. koz- sent to me by him, however, certain additional loffi, this species closely resembles E. pacificus. Both information was revealed by mounting specimens species share the same spination formula with E. pilosus, E. ehlersi, E. worthingi, and E. ferrun- in Hoyer's and observing them with differential gineus (E. dujardinii is similar but has a lateral interference contrast optics. I have reillustrated accessory spine on segment 10). E. pacificus (Figures 26-31) in order to facilitate the comparison of this species with others that Echinoderes ehlersi, E. worthingi, and E. fer- might be confused with it. rungineus are smaller, 210-260 \ua trunk length, although the latter species shares with E. pacificus Segments 3-12 are covered with short hairs in a the presence of a sensory hair posterior to the distinctive pattern (Figures 26, 27). A fine pectinate lateral spine on segment 12. Echinoderes pacificus fringe is evident ventrally on the posterior border is most easily distinguished from all species having of segments 4-13 although it is much less distinct the same spination formula by its short lateral spine on segment 12; segments 6-13 exhibit this fringe on segment 12 and the prominent border of hairs dorsally. on the terminal sternal plates. As noted in the discussion of E. kozloffi, the short (12-17 urn) lateral spine on segment 12 of E. pacificus is diagnostic. This spine tends to curve Echinoderes brevicaudatus, substitute name away from the trunk. Slightly posterior to it, at the for £. brevispinosus Higgins junction of segments 12 and 13, there is a sensory hair (Figures 28-31) not reported in the original During the course of the present investigation, description. I discovered that the name E. brevispinosa was first Small muscle scars (2-3 \im in diameter), similar used by Metschnikoff (1869:190), corrected to E. to sensory spots, are present anterolaterally on the brevispinosus by Panceri (1876:4), and synonomized ventral surface of the first trunk segment. Narrow, with E. dujardinii by Zelinka (1928:228). Since I slitlike scars occur posterior to the sensory spots inadvertently applied this preoccupied name to a on sternal plates 7-9. More distinctive muscle species from the Red Sea (Higgins, 1966a), I now scars (Figure 26) occur near the lateral margin of propose that it be replaced by the substitute name, the sternal plates of segment 9; similar scars are Echinoderes brevicaudatus. 22 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY 100pm 11111111111111 FICURES 26-27.—Echinoderes paciftcus, paratypic female (USNM 5SSS6 RH162.1), neck and trunk segments: 26, ventral view; 27, dorsal view. NUMBER 248 23 50um 31 FIGURES 28-31.—Echxnoderes pactficus, segments 12-13: 28, ventral view, lateral half, paratypic female (USNM 53336, RH162.1); 29, dorsal view, lateral half of same paratypic female; 30, ventral view, lateral half, paratypic male (USNM 53336, RH162.9); 31, dorsal view, lateral half of same paratypic male. Literature Cited Abe, Y. Dujardin, F. 1930. Das Vorkommen von Echinoderes in den japanischen 1851. Sur un petit animal marin, I'Echinodere, formant un Gewassern. Journal of Science of the Hiroshima Uni- type intermediate entre les Crustaces et les Vers. versity, series B, l(l):39-44, 2 figures, 1 plate. Annales des Sciences Naturelles, Zoologie, series 3, Ax, P., and R. Ax 15:158-160, 1 plate. 1967. Turbellaria Proseriata von der Pazifikkuste der USA Gosse, P. H. (Washington), I: Otoplanidae. Zeitschrift fiir Mor- 1864. The Natural History of the Hairy-backed Animal- phologic und Okologie der Tiere, 61:215-254, 18 cules. The Intellectual Observer: Review of Natural figures. History, Microscopic Research, and Recreative 1974. Interstitielle Fauna von Galapagos, V: Otoplanidae Science, 5:387-406, 2 plates. (Turbellaria, Proseriata). Mikrofauna des Meeres- Greeff, R. bodens, 27:1-28, 11 figures. 1869. Untersuchungen • iiber einige merkwiirdige Formen Bacescu, M., £. Dumitrescu, A. Marcus, G. Paladian, and R. des Anthropoden- und Wurm-Typus. Archiv fiir Mayer Naturgeschichte, 35(l):71-100, 4 plates. 1963. Donnces quantitatives sur la faune petricole de la Grobben, K. mer noire a Agigea (Secteur Roumain), dans les 1905. Kinorhyncha. Pages 335-336 in C. Claus and K. conditions speciales de l'annee 1961. Travaux du Grobben, Lehrbuch der Zoologie. x + 955 pages, Museum d'Histoire Naturelle "Grigore Antipa," 996 figures. Marburg: N. G. Elwert'sche Verlagsbuch- 4:131-155, 3 figures, 1 plate. handlung. Blake, C, H. Hartog, M. 1896. Rotifera, Gastrotricha, and Kinorhyncha. Pages 197- 1930. Three New Species of Worms Belonging to the 238 (15 figures) in volume 2 in S. F. Harmer and Order Echinodera. Biological Survey of the Mount A. E. Shipley, editors, The Cambridge Natural His- Desert Region, 4:3-10, 8 figures. tory, xii + 560 pages, 257 figures. London: Mac- Boykin, J. C. Millan and Co., Ltd. 1974. Phylum Kinorhyncha. Pages 39-41 in E. N. Kozloff, Higgins, R. P. Keys to the Marine Invertebrates of Puget Sound, 1960. A New Species of Echinoderes (Kinorhyncha) from the San Juan Archipelago, and Adjacent Regions. Puget Sound. Transactions of the American Micro- vii + 266 pages, 1 unnumbered figure. Seattle: scopical Society, 79(1):85-91, 1 figure. University of Washington Press. 1961. Three New Homalorhage Kinorhynchs from the Cams, J. V. San Juan Archipelago, Washington. Journal of the 1885. Prodromus faunae Mediterraneae. Volume 1, xi + Elisha Mitchell Scientific Society, 77(l):81-88, 14 525 pages. Stuttgart: E. Schweizerbart'sche Verlag- figures. shandlung. 1964a. Redescription of the Kinorhynch Echinoderes Chitwood, B. G. remanei (Blake, 1930) Karling, 1954. Transactions of 1951. Echinoderella steineri, new species (Scolecida, Echi- the American Microscopical Society, 83(2):243-247, nodera). Texas Journal of Science, 3(1): 113-114, 1 3 figures. unnumbered figure. 1964b. Three New Kinorhynchs from the North Carolina 1964a. European Kinorhynchs from Tomales Bay, Cali- Coast. Bulletin of Marine Science of the Gulf and fornia. Abstract 6, pages 2-3 in Abstracts of Contrib- Caribbean, 14(3):479-493, 18 figures. uted Papers, Forty-fifth Annual Meeting of the 1966a. Faunistic Studies in the Red Sea (in Winter, 1961- Western Society of Naturalists. 1962), Part II: Kinorhynchs from the Area of Al- 1964b. The Intertidal Occurrence of Echinoderes pennaki Ghardaqa. Zoologisches Jahrbucher, Systematik, (Kinorhyncha) in the Straits of Juan de Fuca, Wash- Oekologie und Geographic der Tiere, 93:118-126, 9 ington. Abstract 7, page 3 in Abstracts of Contrib- figures. uted Papers, Forty-fifth Annual Meeting of the 1966b. Echinoderes arlis, a New Kinorhynch from the Western Society of Naturalists. Arctic Ocean. Pacific Science, 20(4):518-520, 2 figures. Claparede, E. 1967. The Kinorhyncha of New Caledonia. In Expedition 1863. Beobachtungen iiber Anatomic und Entwickjungs- Frangaise sur Recifs Coralliens de la Nouvelle Cale- geschichte wirbelloser Tiere an der Kuste der Nor- donia, 2:75-90, 12 figures. mandie angestellt. 120 pages, 18 plates. Leipzig: 1968. Taxonomy and Postembryonic Development of the Wilhelm Engelmann. Cryptorhagae, a New Suborder for the Mesopsam- 24 NUMBER 248 25 mic Kinorhynch Genus Cateria. Transactions of the butions du Laboratoire de Zoologie, Academic Na- American Microscopical Society, 87(l):21-39, 25 tionale de Peiping, l(4):l-9, 1 plate. figures. Ludwig, H. 1969a. Indian Ocean Kinorhyncha, 1: Condyloderes and 1886. Dr. Johannes Leunis Synopsis der Tierkunde. Vol- Sphenoderes, New Cyclorhagid Genera. Smithsonian ume 2, 1231 pages, 1160 figures. Hannover: Contributions to Zoology, 14:1-13, 23 figures. Hahn'sche Buchhandlung. 1969b. Indian Ocean Kinorhyncha, 2: Neocentrophyidae, a Marinov, T. New Homalorhagid Family. Proceedings of the Bio- 1964. On the Microzoobenthos Fauna of the Black Sea logical Society of Washington, 87(7): 113-128, 5 fig- (Kinorhyncha and Halacaridae) [in Bulgarian with ures. English summary]. Bulletin of the Institute of Fish 1971. A Historical Overview of Kinorhynch Research. Culture and Fisheries, Varna KH, 4:61-71, 11 figures. Pages 25-31 in N. C. Hulings, editor, Proceedings Mclntyre, A. D. of the First International Conference on Meiofauna. 1962. The Class Kinorhyncha (Echinoderida) in British Smithsonian Contributions to Zoology, 76:1-205, 1 Waters. Journal of the Marine Biological Association figure. of the United Kingdom, 42:503-509, 2 figures. 1974. Kinorhyncha. Chapter 11, pages 507-518 in volume Merriman, J. A., and H. O. Corwin 1 in A. C. Giese and J. S. Pearse, editors, Reproduc- 1973. An Electron Microscopical Examination of Echino- tion of Marine Invertebrates, xi + 546 pages, 18 deres dujardini Claparede (Kinorrhyncha) [sic]. figures. New York: Academic Press. Zeitschrift fur Morphologic der Ticre, 76:227-242, d'Hondt, J. L. 13 figures. 1973. Contribution a l'£tude de la Microfaune Interstiti- Metschnikoff, E. elle des Plages de l'Ouest Alge"rien. Vie et Milieu, 1865. Ueber einige wenig bekannte niedere Thierformen. 23(2):227-241, 2 figures. Zeitschrift fur wissenschaftliche Zoologie, 15:450-461, Hyman, L. H. 1 plate. 1951. The Invertebrates. Volume 3, vi + 572 pages, 223 1869. Bermerkungen iiber Echinoderes: Melanges biolo- figures. New York: McGraw Hill. giques 7. Bulletin de I'Academic des Sciences de Saint Johnston, T. H. Petersbourg, 4:190-194. 1938. Report on the Echinoderida. In Scientific Reports of Mobius, K. the Australasian Antarctic Expedition, 1911-1914, 1884. Nachtrag zu dem im Jahre 1873 erschienenen Ver- series C, 10(7): 1-13, 7 figures. zeichniss der wirbellosen Thiere der Ostsee. Pages Karling, T. G. 1-10 in volume 4 in Bericht der Kommission zur 1954. Echinoderes levanderi n. sp. (Kinorhyncha) aus der wissenschaftlichen Vntersuchung der deutschen Ostsee. Arkiv for Zoologi, series 2, 7(10):189-192, 6 Meere, Berlin. figures. 1887. Systematische Darstellung der Thiere des Plankton Kirsteuer, E. gewonnen in der westlichen Ostsee Fahrt von Kiel 1964. Zur Kenntnis der Kinorhynchen Venezuelas. Zo- in den atlantischen ocean bis jenseit der Hebriden. ologischer Anzeiger, 173(6):388-393, 2 figures. Pages 111-124 in volume 5 in Bericht der Kommis- Kozloff, E. N. sion zur Untersuchung der deutschen Meer, Berlin, i 1972. Some Aspects of Development in Echinoderes (Kino- Moore, P. G. rhyncha). Transactions of the Americal Microscopical 1973. Campyloderes macquariae Johnston, 1938 (Kinorhyn- Society, 91(2):119-130, 18 figures. cha: Cyclorhagida) from the Northern Hemisphere. Lang, K. Journal of Natural History, 7:341-354, 4 figures, 1949. Echinoderida. In N. H. Odhner, editor, Further 1 plate. Zoological Results of the Swedish Antarctic Expedi- Nyholm, K.-G. tion, 1901-1903, 4(2): 1-22, 8 figures. 1947a. Contributions to the Knowledge of the Postembry- Leuckart, R. onic Development in Echinoderida Cyclorhagae. 1854. Bericht iiber die Leistungen in der Naturgeschichte Zoologiska bidrag frdn Uppsala, 25:423-428, 4 figures. der niederen Thiere wahrend der Jahre 1848-1853. 1947b. Studies in the Echinoderida. Arkiv fur Zoologi, Archiv fur Naturgeschichte, 20(2):289-473. 39A(14):l-36, 22 figures, 2 plates. 1869. Bericht iiber die wissenschaftlichen Leistungen in der Omer-Cooper, J. Naturgeschichte der niederen Thiere wahrend der 1957. Deux nouvelles especes de Kinorhyncha en prove- Jahre 1868-1869. Archiv fur Naturgeschichte, 35(2): nance de l'Afrique du Sud. Bulletin M ensue I de la 207-344. Socie'te Linnienne de Lyon, 26(8):213-216, 3 figures. Levander, K. M. Pagenstecher, H. A. 1900. Uber das Herbst- und Winterplankton im finnischen 1875. Echinoderes Sieboldii. Zeitschrift fur wissenschaft- Meerbusen und in der Alands-See, 1898. Acta So- liche Zoologie, supplement, 25(S):117-123, 1 plate. cietatis pro Fauna et Flora Fennica, 18(5): 19-20. 1877. Allgemeine Zoologie. Part 2. vi + 528 pages, 206 Lou, T. H. figures. Berlin: Wiegandt, Hempel & Parey. 1934. Sur la presence d'un nouveau Kinorhynque a Panceri, P. Tchefou: Echinoderes tchefouensis sp. nov. Contri- 1876. Osservazioni intorno a nuove forme di Venni Nema- 26 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY todi marini. Atti delta Reale Accademia delle Southern, R. Scienze Fisiche e Matematiche, 7(10):l-9, 1 plate. 1914. Nemathelmia, Kinorhyncha, and Chaetognatha. Part Perrier, R. 54 in Clare Island Survey. Proceedings of the Royal 1893. Elements d'Anatomie comparee. 1208 pages. Paris: Irish Academy, 31:1-80, 12 plates. Librairie J.-B. Bailliere et Fils. Timm, R. W. Reinhard, W. 1958. Two New Species of Echinoderella (Phylum 1881. Cber Echinoderes und Desmoscolex der Umgegend Kinorhyncha) from the Bay of Bengal. Journal of von Odessa. Zoologischer Anzeiger 4(97):588-592. the Bombay Natural History Society, 55(l):107-109, 1885. Kinorhyncha: Their Anatomical Structure and Posi- 2 figures. tion in the System [in Russian]. 101 pages, 5 plates. Tokioka, T. Kharkov: University Printing Office. 1949. Notes on Echinoderes Found in Japan. Publications 1887. Kinorhyncha (Echinoderes), ihr anatomischer Bau of the Seto Marine Biological Laboratory, l(2):67-69, und ihre Stellung im System. Zeitschrijt fur wissen- schaftliche Zoologie, 45:401-467, 3 plates. 1 figure. Reraane, A. Zaneveld, J. S. 1928. Kinorhyncha. In G. Grimpe and E. Wagler, editors, 1938. Marine Gastrotricha and Kinorhyncha from Sche- Die Tierwelt der Nord- und Ostsee, 7(d,):57-84, 20 veningen. Zoologische Mededeelingen, 20:257-262. figures. Zelinka, C. 1936. Gastrotricha und Kinorhyncha. Number 2 in part 1896. Demonstration von Tafeln der Echinoderes-Mono- 1 in section 2 of volume 4 in H. G. Bronn, editor, graphic Verhandlungen der Deutschen Zoologischen Klassen und Ordnungen des Tierreichs. vi + 243- Gesellschaft, 6:197-199. 385, 297 figures. 1907. Zur Kenntnis der Echinoderen. Zoologischer An- Schepotieff, A. zeiger, 32(5):130-136. 1907a. Zur Systematik der Nematoideen. Zoologischer An- 1908. Zur Anatomie der Echinoderen. Zoologischer An- zeiger, 31(5-6):132-161, 25 figures. zeiger, 33(19-20) :629-647, 11 figures. 1907b. Die Echinoderiden. Zeitschrift fur wissenschaftliche Zoologie, 88(2):291-326, 4 plates. 1912. Die Spermatozoen der Echinoderen und ihre Genese. Schmidt, P. Pages 520-527 in Verhandlungen des VIII Inter- 1974. Interstitielle Fauna von Galapagos, X: Kinorhyncha. nationalen Zoologen-Kongress zu Graz vom 15.-20. Mikrofauna des Meeresbodens, 43:1-15, 2 figures. August 1910. 10 figures. Schultze, M. 1913. Die Echinoderen der Deutschen Sudpolar-Expedition 1853. Ueber Chaetonotus und Ichtydium (Ehrb.) und eine 1901-1903. In Deutschen Sudpolar-Expedition 1901- neue verwandte Gattung Turbanella. Archiv fur 1903, 14(Zoology 6):419-436, 1 plate. Anatomie, Physiologic und wissenschaftliche Medicin. 1928. Monographic der Echinodera. iv + 396 pages, 73 1853:241-254, 1 plate. figures, 27 plates. Leipzig: Wilhelrn Engelmann. REQUIREMENTS FOR SMITHSONIAN SERIES PUBLICATION Manuscripts intended for series publication receive substantive review within their originating Smithsonian museums or offices and are submitted to the Smithsonian Institution Press with approval of the appropriate museum authority on Form SI-36. 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