The Atlantic Barrier Reef Ecosystem at Carrie Bow Cay, Belize, II: Kinorhyncha SERIES PUBLICATIONS OF THE SMITHSONIAN INSTITUTION Emphasis upon publication as a means of "diffusing knowledge" was expressed by the first Secretary of the Smithsonian. In his formal plan for the Institution, Joseph Henry outlined a program that included the following statement: "It is proposed to publish a series of reports, giving an account of the new discoveries in science, and of the changes made from year to year in all branches of knowledge." This theme of basic research has been adhered to through the years by thousands of titles issued in series publications under the Smithsonian imprint, commencing with Smithsonian Contributions to Know/edge in 1848 and continuing with the following active series: Smithsonian Contributions to Anthropology Smithsonian Contributions to Astrophysics Smithsonian Contributions to Botany Smithsonian Contributions to the Earth Sciences Smithsonian Contributions to the Marine Sciences Smithsonian Contributions to Paleobiology Smithsonian Contributions to Zoology Smithsonian Studies in Air and Space Smithsonian Studies in History and Technology In these series, the Institution publishes small papers and full-scale monographs that report the research and collections of its various museums and bureaux or of professional colleagues in the world of science and scholarship. The publications are distributed by mailing lists to libraries, universities, and similar institutions throughout the world. Papers or monographs submitted for series publication are received by the Smithsonian Institution Press, subject to its own review for format and style, only through departments of the various Smithsonian museums or bureaux, where the manuscripts are given substantive review. Press requirements for manuscript and art preparation are outlined on the inside back cover. S. Dillon Ripley Secretary Smithsonian Institution SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES •NUMBER 18 The Atlantic Barrier Reef Ecosystem at Carrie Bow Cay, Belize, II: Kinorhyncha Robert P. Higgins SMITHSONIAN INSTITUTION PRESS City of Washington 1983 ABSTRACT Higgins, Robert P. The Atlantic Barrier Reef Ecosystem at Carrie Bow Cay, Belize, II: Kinorhyncha. Smithsonian Contributions to the Marine Sciences, number 18, 131 pages, 343 figures, 23 tables, 1983.—Eighteen new species, including one new genus of Kinorhyncha, are described from the reef ecosystem at Carrie Bow Cay, Belize. Pycnophyes neapolitanus Bacescu, 1968, is considered a junior synonym of P. ponticus Zelinka. Pycnophyes quadndentatus Zelinka, 1928, and P. flagellatus Zelinka, 1928, are synonymized under the former taxon and placed in Paracentrophyes, new genus (Neocentrophyidae), represented by a new species from the study area. Other genera represented by the remaining new species are extensively reviewed, species names are corrected to agree in gender, distribution records and keys to adults are compiled, and phylogeny discussed. Species distribution and richness are discussed. A maximum of 13 species representing four genera were found in a single local sample. This is contrasted with similar data from other parts of the world. Certain local species appeared to prefer or were restricted to fine, organically rich, low energy mangrove sediments as opposed to the more heterogeneous sediments with Thalassia beds and the even higher energy sediments of the coral reef proper. OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: Seascape along the Atlantic coast of eastern North America. Library of Congress Cataloging in Publication Data Higgins, Robert P. The Atlantic Barrier Reef ecosystem at Carrie Bow Cay, Belize, II—Kinorhyncha. (Smithsonian contributions to the marine sciences ; no. 18) Bibliography: p. Supt.ofDocs.no.: SI 1.41:18 1. Kinorhyncha—Belize—Carrie Bow Cay. 2. Coral reef ecology—Belize—Carrie Bow Cay. 3. Carrie Bow Cay (Belize) I. Title. II. Series. QL391.K5H53 1983 574.5'26367'097282 82-600235 Contents Page Introduction 1 Methods 4 Acknowledgements 7 Order CYCLORHAGIDA Zelinka, 1896 7 Family ECHINODERIDAE Biitschli, 1876 7 Genus Echinoderes Claparede, 1863 7 Key to Adults of Echinoderes 7 Echinoderes abbreviatus, new species 10 Echinoderes horni, new species 15 Echinoderes imperforatus, new species 20 Echinoderes truncatus, new species 26 Echinoderes wallaceae, new species 31 Discussion of Echinoderes 36 Order HOMALORHAGIDA Zelinka, 1896 43 Family NEOCENTROPHYIDAE Higgins, 1969b 43 Paracentrophyes, new genus 44 Paracentrophyes praedictus, new species 44 Family PYCNOPHYIDAE Zelinka, 1896 54 Genus Pycnophyes Zelinka, 1907 54 Key to Adults of Pycnophyes 54 Pycnophyes corrugatus, new species 56 Pycnophyes ecphantor, new species 60 Pycnophyes emarginatus, new species 64 Pycnophyes iniorhaptus, new species 68 Pycnophyes longicornis, new species 72 Discussion of Pycnophyes 76 Genus Kinorhynchus Sheremetevskij, 1974 81 Key to Adults of Kinorhynchus 81 Kinorhynchus apotomus, new species 82 Kinorhynchus belizensis, new species 85 Kinorhynchus deirophorus, new species 89 Kinorhynchus distentus, new species 93 Kinorhynchus erismatus, new species 98 Kinorhynchus stenopygus, new species 102 Kinorhynchus trisetosus, new species 108 Discussion of Kinorhynchus 115 Species Distribution and Richness 116 Phylogenetic Relationships 126 Literature Cited 128 The Atlantic Barrier Reef Ecosystem at Carrie Bow Cay, Belize, II: Kinorhyncha Robert P. Higgins Introduction Since their discovery in 1841 (Dujardin 1851), there have been 119 reports on the distribution of the Kinorhyncha, a phylum of marine meio- benthic invertebrates. Only a few dozen of these reports have involved tropical shallow-water eco- systems. Five reports have been based on studies of coral reef ecosystems. The most extensive of these stud- ies was conducted on the coral reefs of Baie St. Vincent, New Caledonia (Higgins, 1967). These resulted in the description of two new species, Echinoderes newcaledoniensis Higgins, 1967, and Sem- noderes pacificus Higgins, 1967, and a new distri- bution record for Campyloderes macquariae John- ston, 1938, which was known originally only from Macquarie Island, in polar seas to the south. Three additional species have been reported from coralline sediments of the Red Sea; these include Echinoderes riedli Higgins, 1966a, E. brevicaudatus Higgins, 1966a, and Pycnophyes egyptensis Higgins, 1966a. Centroderes spinosus (Reinhard, 1881) has been found in the coralline sandy silt of Castle Harbor, Bermuda (Coull, 1968, 1970), and Neo- centrophyes intermedius Higgins, 1969b, was reported from fine carbonate sediments of Nosy Be, Mad- agasgar. Robert P. Higgins, Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. Mangrove sediments, normally consisting of a fine mud, are a productive tropical shallow-water habitat for kinorhynchs. Kirsteuer (1964) was the first to make such a report in which he described Echinoderes caribiensis Kirsteuer, 1964, from the Bay of Mochima, Venezuela. Several years later, I described Sphenoderes indicus Higgins, 1969a, from mangrove mud in the Bay of Kutch, near Jamnigar, India. Intertidal and subtidal sediments, not specifi- cally identified as coralline in origin or associated with coral reef ecosystems, have accounted for the remaining tropical shallow-water kinorhynch dis- tribution records. From the interstitial habitat of high energy sand beaches or subtidal sand, three species of Catena have been described. Two of these species are restricted to tropical intertidal beaches. The first species, Catena styx Gerlach, 1956, was described from Brazil and reported from Angola, on the opposite side of the South Atlantic, a year later (Delamare-Deboutteville, 1957). Since then it has been recorded from both the original locality and a second locality in Brazil (Higgins, 1968) as well as from several localities on the east coast of India (Ganapati and Rao, 1962; Rao and Ganapati, 1966, 1968; Nagabhu- shanam, 1972). A second species, Catena gerlachi Higgins, 1968, was described from the east coast of India, reported again from the same locality by Rao and Ganapati (1968), and more recently reported from the Andaman Islands, on the op- SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES posite side of the Bay of Bengal (Higgins and Rao, 1979). Echinoderes pacificus Schmidt, 1974, was reported from subtidal heterogenous sediments of the Gal- apagos Islands in the Pacific Ocean; all other records are from the Indian Ocean. Echinoderes bengalensis (Timm, 1958) was described from sub- tidal sand at Sonadia Island, near Cox Bazar, Bangladesh. A second species, E. sonadiae (Timm, 1958), was described from the same collection but has been synonymized with E. bengalensis (Hig- gins, 1977b) as a juvenile stage of the latter species. This species has also been found along with Catena styx and C gerlachi on the east coast of India (Rao and Ganapati, 1968). Condyloderes paradoxus Higgins, 1969a, Sphenoderes indicus Hig- gins, 1969a, and Neocentrophyes satyai Higgins, 1969b, were described from the east coast of India. Damodaran (1972) reported an unidenti- fied species of Echinoderes from the southwest coast of India. Echinoderes andamanensis Higgins and Rao, 1979, Echinoderes ehlersi Zelinka, 1913 (orig- inally found in sediment washed from an uniden- tified invertebrate collected in Zanzibar), and one species of Pycnophyes have been reported from the Andaman Islands (Higgins and Rao, 1979). Ki- norhynchs have been collected from the Arabian Gulf near Bahrain (Basson et al., 1977) and the estuarine sediments of Shott-al-Arab, Iraq (Saad and Arlt, 1977), but were not identified further. In April, 1977, I collected kinorhynchs in con- junction with the Smithsonian Institution's long- term study of the coral reef ecosystem at Carrie Bow Cay, Belize (Figure 1). Carrie Bow Cay (16°48.1'N, 88°04.7'W) is a small island, one of many that exist as part of the Belizian Barrier Reef which extends in a north-south direction and separates an extensive lagoon system from the open water of the Caribbean Sea. Two hundred meters north of Carrie Bow Cay, begin- ning at the edge of a Thalassia seagrass commu- nity in the shallowest part of the lagoon (1.5-2 m), is a study transect 50 m wide running east- ward, 650 m to the open ocean. The lagoon area extends westward 20 km at Carrie Bow Cay. The western portion is as much as 20 m deep, but near the barrier reef, it rarely exceeds 5 m. Shal- low seagrass bottoms {Thalassia testudinium), patch reefs, and mangrove cays are predominant on the outer barrier platform. A more complete descrip- tion of this ecosystem has been compiled by Riit- zler and Maclntyre (1982). For purposes of sampling in as many repre- sentative sediment types as practical within this ecosystem, the first station (RH 442) was estab- lished inside the shallow channel separating the two component mangrove cays, 2 km northwest of Carrie Bow Cay, which give Twin Cays (16°50.0'N, 88°06.0'W) its name. The meander- ing channel, especially near its southwestern lim- its, attains its maximum depth of 1-2 m where a fine gray calcareous sediment mixed with man- grove and seagrass detritus and sand accumu- lates. It represents the most minimal energy hab- itat of the series of stations established in this study. The second station (RH 443) was 500 m south- west of the first, at a depth of 3.3 m. Here the bottom consisted of some scattered seagrass in a sediment made up of coralline mud mixed with sand and seagrass detritus. The sediment was detectably more coarse than the first station, suggesting a higher energy system. The third station (RH 444) was located 500 m south of the previous station. The water depth was 3.0 m, the seagrass was more abundant, and the sediment, although similar to the previous station, was finer, suggesting a lesser energy level, possibly related to the increased amount of sea- grass at this site. The last three stations were established on the study transect, beginning with RH 446 at or near the zero point of the transect, 200 m north of Carrie Bow Cay. This station was located at a depth of 2.0 m in the Thalassia zone where sedi- ment was mixed Halimeda sand and shell with some fine material. The second transect station (RH 447) was established in the sand and rubble zone of the lagoon, 1.5 m deep. Here the sediment consisted of Halimeda sand with broken shell and very little fine sediment. No areas of suitable sediments existed between this latter station and NUMBER 18 South Water Cay Carrie Bow Cay 17 N 443® 444 (§) 88 W Twin Cays 445 (§) FIGURE 1.—Map of the study sites, Twin Cays and Carrie Bow Cay, Belize, showing the location of the six sampling stations (RH 442-447); area of larger map located on inset by SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES the sand trough station (RH 445) located at a depth of 25 m where very fine sand accumulated. Although the Thalassia zone probably repre- sented the lowest energy system within the study transect, all three stations were obviously sub- jected to periodically if not constantly higher levels of wave energy than the three stations 2 km further back in the lagoon. METHODS.—In order to maximize the amount of kinorhynchs collected, a meiobenthic dredge was used to collect the surface few centimeters of sediment at each station. Insofar as it was possi- ble, each sample consisted of a similar amount of sediment, about 10 liters, which probably repre- sents a 4 m2 minimum sample surface area, as- suming the 25 cm wide dredge blade removed only the upper centimeter of sediment. Though by no means a quantitative method, samples taken in this manner probably offer a reasonable estimate of the relative abundance of kinorhynchs and their species diversity at a given station. In the laboratory, sea water was added to each container of sediment in amounts sufficient to facilitate adequate mixing to produce a suspen- sion into which fine air bubbles were introduced. Kinorhynchs have a hydrophobic exoskeleton which, as in several other meiobenthic taxa such as amphipods, taniads, nematodes, ostracodes, etc., transports the animals to the surface film where they remain trapped by the surface tension. After a few moments during which the suspended sediments settled away from the surface film, a half-sheet of standard commercial white inexpen- sive bond paper was allowed to lightly touch the surface film and was withdrawn quickly, thereby removing the animals trapped in the surface film. The attached meiobenthic organisms, mostly kinorhynchs and amphipods in this instance, were then washed off the paper onto a 62 /mi mesh screen, and the bubbling process repeated until no further specimens were collected. Specimens were preserved in 5 percent for- malin and then transferred to 70 percent ethyl alcohol. A few specimens of what, under low magnification, appeared to be the most common species, were removed for SEM study; the re- mainder were transferred to a 70 percent ethyl alcohol-5 percent glycerin solution that was al- lowed to evaporate to glycerin. Most specimens were removed from the glycerin and individually placed in a modified Hoyer's mounting medium, between two coverslips, and positioned on Cobb aluminum slide frames. This mounting procedure allows the slide to be placed on either of its surfaces so that both dorsal and ventral aspects of the specimen can be observed. Hoyer's medium is necessary to soften the spec- imen so that, by judicious manipulation of the coverslip, the specimen will assume a dorsoventral position; this medium also clears the specimen, thus revealing the detailed structure of the exo- skeleton. A disadvantage of normal Hoyer's medium is its tendency to clear the specimen too much, especially over a period of several years. This may be partially overcome by using only 125 grams of chloral hydrate instead of the 200 grams normally required. Coverslips were sealed with an epoxy paint. Each specimen was studied with the use of Zeiss differential interference contrast optics and analyzed. The resulting data are expressed in a standard format of abbreviations and terminol- ogy (Higgins, 1967, 1969a). Measurements are given in micrometers (jum); ratios (i.e., SW/TL) are expressed in percent of the total length (TL) measured on the midline, from the anterior mar- gin of segment 3 (first trunk segment) to the posterior margin of segment 13, exclusive of spines. Maximum sternal width (MSW) is meas- ured at the anteroventral margin of the widest pair of sternal plates as first encountered in meas- uring each segment from anterior to posterior. Sternal width at segment 12 (SW), or standard width, is measured at the anteroventral margin of the 12th sternal plates. Placids (neck plates of segment 2) and, where applicable, trichoscalid plates of cyclorhagid taxa are numbered begin- ning with the midventral placid as zero; therefore, those on either side of the midventral placid are each number 1, those next in sequence, number 2, etc. NUMBER 18 Oral Style Mouth Cone Spinoscalid Trichoscalid Plate Trichoscalid Placid Sensory Spot Pectinate Fringe Lateral Spine (L-4) Adhesive Tube Maximum Sternal Width (MSW) Lateral Accessory Spine (LA-10) L-10 Standard Width (SW) L-11 Gonopore -12 ■13 Lateral Terminal Accessory Spine Lateral Terminal Spine Penile Spine (P-1) P-2 P-3 L-7 L-8 L-9 Pachycyclus Muscle Scar • o cf a»o 0 r„» Cuticular Scar Middorsal Spine (D"6) 8#syy Perforation Sites —:''.i 50pm ".""J'»/,/ MM /\^vjY< ;;-..:! * I* SP 170 171 NUMBER 18 71 TABLE 12.- -Measurements (/xm) and indices (%) for Pycnophyes imorhaplus adults (see "Methods' for character abbreviations) Cha.ra.ctcr Standard Standard Coefficient Number Range Mean deviation 39.1 error 14.8 of variability TL 6 7 572-686 631.4 6.2 2 2 634-676 655.0 29.7 21.0 4.5 62 9 572-686 636.7 36.9 12.3 5.8 SW 6 7 120-132 126.3 4.5 1.7 3.6 2 2 140-144 142.0 2.8 2.0 2.0 62 9 120-144 129.8 8.0 2.7 6.2 SW/TL 6 7 17.8-21.6 20.1 1.4 0.5 7.0 2 2 20.7-22.7 21.7 1.4 1.0 6.5 62 9 17.8-22.7 20.4 1.5 0.5 7.4 MSW-6 6 7 144-152 148.0 4.0 1.5 2.7 2 2 152-156 154.0 2.8 2.0 1.8 62 9 144-156 149.3 4.5 1.5 3.0 MSW/TL 6 7 21.0-24.8 23.5 1.5 0.6 6.2 2 2 22.4-24.6 23.5 1.6 1.1 6.6 62 9 21.0-24.8 23.5 1.4 0.5 5.8 LTS 6 7 152-180 170.3 9.2 3.5 5.4 2 2 108-112 110.0 2.8 2.0 2.6 62 9 108-180 156.9 27.8 9.3 17.7 LTS/TL 6 7 24.4-29.3 27.1 2.3 0.9 8.4 2 2 16.0-17.7 16.9 1.2 0.9 7.1 62 9 16.0-29.3 24.8 4.9 1.7 19.9 166); anterior margin of tergal plate of segment even, line of punctate sculpturing parallel to mar- gin; episternal plates with small area of thin cuticle at anterior margin, no such area on mid- sternal plate; midsternal plate trapezoidal, about 83 /xm long, 60 /xm basal width with uneven margin tapering to 30 /xm at apex (Figures 161, 165). Sternal plate 4 of male with prominent adhe- sive tubes; about 40 /xm long (Figure 161), no setae near lateral margin but present in female (Figure 165); areas of punctate cuticular sculp- turing already noted previously. Segments 5-11 FIGURES 169-174.—Pycnophyes iniorhaptus, new species: 169, segments 2-4, ventral view; 170, segments 7, 8, ventral view; 171, segments 11-13, ventral view; 172, segments 2-4, dorsal view; 173, segments 7, 8, dorsal view; 174, segments 11-13, dorsal view. (Interference contrast photographs all with same scale (in jum) as shown in Figure 169; SP = spermatozoa.) similar; segment 12 with margin of sternal plates extending slightly posterior at the ventral mid- line; tergal margin even. Posterior margin of each sternal plate of terminal segment angular (Figure 163) at least in males, structure not clearly evident in females. Tergal margin rounded with slightly fringed edge. Two pairs of penile spines at an- terolateral margins of 12th sternal plates; single gonopore in female. Lateral terminal spines long, 152-180 /xm, 24.4-29.3 percent of trunk length in males, slightly shorter, 108-112 /xm, 16.0-17.7 percent of trunk length in females, slightly re- curved with transverse mark about one-third dis- tance from base (Figures 159, 160). Males differ from females by presence of ad- hesive tubes and lack of setae on segment 4 (Figure 161), 2 pairs of penile spines at anterolat- eral margins of terminal sternal plates (Figure 163), no subdorsal setae, shorter lateral terminal 72 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES spines, slight differences in number and position of cuticular markings, scars, etc., and a slightly denticulate anterodorsal margin on the first trunk segment. Morphometric data for adult specimens are shown in Table 12. HOLOTYPE.—Adult male, TL 676 /xm (Figures 159-164, 169-174), Twin Cays, sta RH 442, Be- lize (16°50.0'N, 88°06.0'W), 8 Apr 1977, col. R.P. Higgins, USNM 70000. ALLOTYPE.—Adult female TL 676 /xm (Figures 165-168), other data as for holotype, USNM 70001. PARATYPES.—Six males and 1 female, TL 572- 686 /xm, other data as for holotype, USNM 70002. REMARKS.—The presence of the areas of dis- tinctive punctate cuticle located laterally on each sternal plate of segments 4-12 of Pycnophyes inio- rhaptus is unique to this genus or any other kino- rhynch taxon. A second important character is the line of cuticular punctations near the anterior tergal margin of the first trunk segment. Most sculpturing of this margin involves a toothed or toothlike pattern restricted to the anteriormost edge without a second pattern more posterior as in Pycnophyes emarginatus or P. sanjuanensis. Others, such as P. corrugatus, have various geometrically arranged cuticular ridges near the lateral edges of the sternal plates. Pycnophyes egyptensis has a band of punctate cuticle on either side of the dorsal midline of the first trunk segment. Pycnophyes egyptensis is similar to P. imorhaptus in that the former species has a similar shaped mid- sternal plate but with a thin area of cuticle near its anterior margins. Pycnophyes egyptensis also has a similar trunk shape, long lateroterminal spines, and distinctive midventral thickenings on seg- ments 11 and 12 only; however, its lateral termi- nal spines do not have the transverse mark, nor are they slightly recurved; it has middorsal setae on most trunk segments and lacks the distinctive areas of punctate cuticle on the sternal plates. ETYMOLOGY.—This species name is from the Greek inion (nape of neck) plus rhaptos (stitched), referring to the transverse line of cuticular punc- tations near the anterior margin of the first trunk segment. Pycnophyes longicornis, new species FIGURES 175-190 DIAGNOSIS.—Trunk tapering slightly beginning with segment 7; lateral terminal spines long, re- curved at tip which is defined by transverse mark about three-fourths distance from base, lateral terminal spines usually perpendicular to trunk upon fixation; middorsal spinous processes at the posterior tergal margins; anterior margin of tergal plate of first trunk segment denticulate; lateral setae on every other segment beginning with segment 4; adhesive tubes absent in males. DESCRIPTION.—Adults (Figures 175-190), trunk length 478-572 /xm, trunk segments tapering slightly in posterior progression beginning with segment 7; MSW-7 120-140 /xm, 23.7-28.5 per- cent of trunk length; SW 108-125 /xm, 20.8-25.4 percent of trunk length. Second segment with 4 dorsal and 4 ventral, even placids. Trunk segments (Figures 175, 176) with some middorsal setae present on females, absent on males, dorsolateral setae, 20/xm long, on segments 3-11, lateral setae at margin of segments 4, 6, 8, 10, with 2 on segment 12, ventrolateral setae on segments 3, 4, 10 with mesial setae on segments 5-9, 11; sensory spots dorsolateral on segments 3- 10, 12, double, on 11, double ventrolateral on segment 4 only, ventrolateral on segments 5-11. Posterior margins of tergal plates 4-11 with prom- inent middorsal spinose processes. Pachycycli well developed; midventral thickenings on segments 10-12 (Figures 175, 187). Muscle scars prominent dorsolaterally and ventrolateral^ on first trunk segment (Figures 175, 176, 185, 188), inverted V- shaped scar especially noticeable near anterome- sial area of thin cuticle on episternal plate of first trunk segment in males (Figure 177); elongate cuticular scars on tergal plates 4-12 transversely oriented in anterior segments, becoming more angular posteriorly, those on ventral plates oval to crescentic, situated mesial to sensory spots (Figures 175, 176). First trunk segment (segment 3) with promi- nent hornlike extensions of the anterolateral mar- gins of tergal plate (Figures, 175, 176); anterior margin of segment denticulate (Figures 178, 182), undulant median ridge of cuticle about one-third NUMBER 18 73 179 180 50pm FIGURES 175-184.—Pycnophyes longicornis, new species: 175, trunk segments, ventral view, holo- typic male (USNM 70003); 176, same, dorsal view; 177, same, segments 3, 4, lateral half, ventral view; 178, same, dorsal view; 179, same, segments 12, 13, lateral half, ventral view; 180, same, dorsal view; 181, segments 3, 4, lateral half, ventral view, allotypic female (USNM 70004); 182, same, dorsal view; 183, same, segments 12, 13, lateral half, ventral view; 184, same, dorsal view. 74 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES [ \ \ I I 187 190 NUMBER 18 75 TABLE 13.—Measurements (jxm) and indices (%) for Pycnophyes longicornis adults (see "Methods" for character abbreviations) C^h si Vs\cXf*r Standard Standard Coefficient V Jlltll Cl^L^ Number Range Mean deviation error of variability TL c? 19 478-551 511.1 20.8 4.8 4.07 2 15 489-572 522.7 26.6 6.9 5.1 62 34 478-572 516.2 23.9 4.1 4.6 SW 6 19 108-124 116.6 4.1 0.9 3.5 2 15 120-125 122.2 2.1 0.6 1.8 62 34 108-125 119.1 4.3 0.7 3.6 SW/TL 6 19 20.8-24.3 22.8 1.2 0.3 5.1 2 15 21.6-25.4 23.4 1.3 0.3 5.4 62 34 20.8-25.4 23.1 1.2 0.2 5.3 MSW-7 6 19 128-136 132.5 2.3 0.5 1.7 2 15 120-140 133.2 4.4 1.1 3.3 62 34 120-140 132.8 3.3 0.6 2.5 MSW/TL 6 19 24.2-28.5 26.0 1.25 0.3 4.8 2 15 23.7-27.1 25.6 1.4 0.4 5.3 62 34 23.7-28.5 25.8 1.3 0.2 5.0 LTS 6 18 148-188 171.3 10.7 2.5 6.2 2 15 140-176 160.3 8.5 2.2 5.3 62 33 140-188 166.3 11.1 1.9 6.7 LTS/TL 6 18 29.4-37.7 33.6 2.2 0.5 6.6 2 15 25.9-34.4 30.8 2.3 0.6 7.3 62 33 25.9-37.7 32.3 2.6 0.5 8.1 distance posterior (Figure 176); episternal plates with small area of thin, wrinkled cuticle at anter- omesial margin, no such area on midsternal plate; midventral plate trapezoidal, about 70 /xm long, 62 /xm basal width with anterior margin slightly expanded, 30 /xm wide at apex (Figure 175). Segment 4 of male similar to that of female (Figures 177, 178, 181, 182), adhesive tubes ab- sent in both sexes, with middorsal seta in female, absent in male, each sternal plate with ventrolat- eral setae and 2 mesial sensory spots; tergal plate with lateral setae and subdorsal sensory spots. Segments 5-9, 11 similar except lateral setae FIGURES 185-190.—Pycnophyes longicornis, new species: 185, segments 3, 4, ventral view; 186, segments 7, 8, ventral view; 187, segments 11-13, ventral view; 188, segments 3, 4, dorsal view; 189, segments 7, 8, dorsal view; 190, segments 11-13, dorsal view. (Interference contrast photographs all with same scale (in /im) as shown in Figure 185.) on segments 6 and 8. Setae on sternal plates 5-9, 11 ventromesial (mesial to sensory spot, Figure 175); seta on sternal plates 10 ventrolateral (lat- eral to sensory spot, Figure 175). Segment 11 with 2 dorsolateral sensory spots on each side. Segment 12 without lateroventral sensory spots or setae; 2 lateral setae and prominent dorsolateral sensory spots present; posterior margin of sternal plates subacute at midline. Terminal segment rounded with slightly angular lateroterminal margins, 2 pairs of penile spines at anterolateral margins of sternal plates, simple gonopores in females. Lateral terminal spines long, 148-188 /xm, 29.4-37.7 percent of trunk length in males, 140- 176 /xm in females; usually perpendicular to trunk upon fixation and with a slight bend near trans- verse mark about 3/4 distance from base. Males differ from females by the presence of 2 pairs of penile spines at anterolateral margins of terminal sternal plates (Figure 179); lack of mid- 76 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES dorsal setae and slightly smaller and different shaped penultimate segment. Morphometric data for adult specimens are shown in Table 13. HOLOTYPE.—Adult male, TL 530 /xm (Figures 175-180, 185-190), Twin Cays, sta RH 442, Be- lize (16°50.0'N, 88°06.0'W), 8 Apr 1977, col. R.P. Higgins, USNM 70003. ALLOTYPE.—Adult female, TL 572 /xm (Figures 181-184), other data as for holotype, USNM 70004. PARATYPES.—Nine females and 9 males, TL 489-556 /xm, other data as for holotype, USNM 70005; 4 females and 2 males, TL 489-551 /xm, Twin Cays, sta RH 443, Belize, other data as for holotype, USNM 70006; 7 males and 1 female, TL 478-530 /xm, Twin Cays, sta RH 444, Belize, other data as for holotype, USNM 70007. REMARKS.—Of the five new species of Pycno- phyes described in this paper, P. longicornis has the broadest relative trunk length; this is especially noticeable when the first trunk segments are com- pared. This new species is similar to P. zelinkaei and P. odhneri in number and position of lateral setae but differs significantly in the number and arrangement of other setae. Like P. zelinkaei, its lateral terminal spines may tend to fix perpendic- ular to the trunk. This assumes that the original illustration of P. zelinkaei was typical of this species as well as accurate. In addition, both species have a dentate anterior margin on the tergal plate of the first trunk segment. In P. zelinkaei this border appeared much different to Zelinka (1928, fig. 64) than to Southern (1914, fig. 33E). The former author's illustration would suggest a row of lon- gitudinal bars forming the dentate appearance. In P. longicornis this area is clearly denticulate, not the same as in P. zelinkaei. The absence of adhesive tubes on the sternal plates of segment 4 in P. longicornis males is similar to P. ecphantor and P. egyptensis. Pycnophyes longicor- nis has a trunk shape similar to the latter species but has very few other similarities. ETYMOLOGY.—This species name is from the Latin longus (long) plus cornus (horn), referring to the lateral terminal spines which, as typically extended, suggest the appearance of the "longhorn" cattle of the southwestern United States. Discussion of Pycnophyes Until 1896, all kinorhynchs were assigned to a single family, Echinoderidae Butschli, 1876, with its monotypic genus, Echinoderes Claparede, 1863. Between 1888 and 1908, Zelinka published a series of papers that were "previews" of his mon- ograph on kinorhynchs (Zelinka, 1928). Zelinka's first comments on kinorhynch classification were directed at a previous attempt (Reinhard, 1885) to divide the Kinorhyncha into three groups of undesignated rank—there is no basis for assuming that these were generic names as stated in a recent paper by Sheremetevskij (1974). In short, Zelinka (1894) stated that "a division of the Echinoderes into Bicerca, Monocerca and Acerca is untenable; the essential reason for dividing the species must be based on the closing apparatus of the anterior end." Thus, he laid to rest Reinhard's artificial divisions that were based on the spine character of the terminal segment. In 1896, Zelinka (1896:198 not 1894 as cited by Sheremetevskij, 1974) erected the family Pycnophyidae but did not establish the genus Pycnophyes until 11 years later (Zelinka, 1907). The first species assigned to this genus was Pycnophyes communis Zelinka, 1908. Although this latter species was not described by Zelinka until 1928, these rules for "indication" were satisfied by the earlier paper. In 1928, Zelinka described 10 new species of Pycnophyes: P. quadndentatus, P. flagellatus, P. echi- noderoides, P. rugosus, P. robustus, P. calmani, P. flav- eolatus, P. communis, P. carinatus, and P. kielensis. Pycnophyes calmani was, in fact, described earlier by Southern, 1914. In addition, Zelinka reas- signed Echinoderes dentatus Reinhard, 1881, and E. ponticus Reinhard, 1881, to the genus Pycnophyes. With the addition of P. zelinkaei Southern, 1914, the number of species assigned to Pycnophyes to- taled 12. At this same time, Zelinka (1928) de- scribed larval species, life history stages known to be Pycnophyes but assigned to new larval genera. NUMBER 18 77 These included Hyalophyes longisetosus, H. solidus, H. conspicuus, Centrophyes curvatus, C. rectilineatus, C. moderatus, C. longihastatus, C. denticulatus, C. diffusus, C. validus, C tenuis, and C. biserratus. Thus, the genus Pycnophyes actually had a total of 24 nomi- nal species. Hyalophyes Zelinka (1907) consisted of the last three stages prior to the adult, and, aside from having a thin cuticle and no secondary sex char- acters, species assigned to this genus were similar to Pycnophyes. Centrophyes Zelinka (1907) consisted of the three stages that precede the Hyalophyes stages. Although they possessed the precursors of the lateral terminal spines, the presence of a midterminal spine dominated their morphology. The larval genus Leptodemus Zelinka, 1907, was assigned to the family Trachydemidae Zelinka (1896), as was the nominal genus Trachydemus (now Kinorhynchus Sheremetevskij, 1974). In his monograph, Zelinka described L. serratus, L. for- ceps, L. forficulus, L. perlatus, L. vitreus, and L. naviculus in addition to reassigning Echinoderes acercus Reinhard, 1881 (considered "unidenti- fiable" by both Zelinka, 1928:360, and Remane, 1936:362), and E. dubius Reinhard, 1887, to this latter genus. These eight species, based on juve- niles lacking all caudal spines (= Acerca sensu Reinhard, 1885) were subject to reinterpretation first by Remane (1929) who stated that L. na- viculus, L. perlatus, L. forficulus, L. dubius, and L. vitreus were members of the genus Pycnophyes and later by Bacescu, 1968, who added L. metschnikowii and L. acercus to this same genus. Two remaining species assigned to Leptodemus, L. forceps and L. serratus, appear to have been ignored. Despite the report of Nyholm (1947b) that Leptodemus can change into a Centrophyes, therefore implying that both the latter two genera are synonyms of Pycnophyes, there is contradictory evidence (Higgins, 1974) that Leptodemus stages are, in fact, juveniles of Kinorhynchus and that the Zelinkian species of Leptodemus are, at best, species incertae sedis. Since 1928, nine additional species of Pycnophyes have been described. These include P. beaufortensis Higgins, 1964b, P. chiliensis Lang, 1953, P. tensis Higgins, 1966a, P. frequens Blake, 1930, P. maximus Reimer, 1963, P. neapolitanus Bacescu, 1968, P. odhneri Lang, 1949, P. sanjuanensis Hig- gins, 1961, and P. sculptus Lang, 1949. In 1963, Hyalophyes pellucidus was synonymized (Reimer, 1963:443) with P. kielensis and sup- ported by Bacescu (1968:243). The latter author, in the same publication, synonymized P. ponticus sensu Bacescu and Bacescu, 1956 (partim) and P. TABLE 14.—Composition of the genus Pycnophyes incorporat- ing the criteria of Zelinka, 1928, by which generic junior synonyms have arisen (asterisk denotes species indeterminata based on inadequate description of the adult) ADULTS JUVENILES Pycnophyes (= Hyalophyes) (= Centrophyes) Lateral terminal spines only Midterminal spines beaufortensis conspicuus bisseratus calmani longisetosus curvatus carinatus solidus denticulatus chiliensis diffusus communis longihastatus corrugatus, moderatus new species dentatus rectilineatus * echinoderoides tenuis ecphantor, validus new species egyptensis emarginatus, new species flaveolatus frequens iniorhaptus, new species kielensis longicornis, new species maximus odhneri 3 ponticus robustus rugosus sanjuanensis sculptus zelinkaei 1= P. communis according to Nyholm (1947b: 10) and Lang (1949:12). 2 Includes Hyalophyes pellucidus. 3 Includes P. neapolitanus. 78 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES TABLE 15.—Distribution of Pycnophyes (see Figure 191; type-localities are in italics; asterisk denotes species indeterminata based on inadequate description of the adult) Species FAO sea area Locality Authority beaufortensis ANW Beaufort (N.C.), USA Higgins, 1964b ASW Bears Bluff (S.C.), USA Higgins, 1964b ASW Marineland (Fla.), USA Higgins, 1964b biserratus ANE W. Sweden Nyholm, 1947b MED Naples, Italy Zelinka, 1928 MED Sukhumi, USSR Sheremetevskij, 1974 calmani ANE Clew Bay, Ireland Southern, 1914 ANE St. Andrews Bay, UK Zelinka, 1928 ANE Oresund, Denmark Lang, 1936 carinatus MED Trieste, Italy Zelinka, 1928 MED Naples, Italy Zelinka, 1928 chiliensis PSW Ouellin Is., Chile Lang, 1953 communis ANE Gullmarfjord, Sweden Nyholm, 1947b MED Naples, Italy Zelinka, 1928 MED Trieste, Italy Zelinka, 1928 conspicuus MED Naples, Italy Zelinka, 1928 MED Trieste, Italy Zelinka, 1928 corrigatus, ASW Twin Cays, Belize Higgins, this study new species curvatus ANE W. Sweden MED Naples, Italy Zelinka, 1928 MED Trieste, Italy Zelinka, 1928 denlalus ANE Clew Bay, Ireland Southern, 1914 ANE Kiel, FRG Zelinka, 1928 ANE Scheveningen, Netherlands Zaneveld, 1938 ANE SW Isle of Man, UK Bruce et al., 1963 MED Odessa, USSR Reinhard, 1881, 1885, 1887, Bacescu, 1968 MED Rumania Bacescu and Bacescu, 1956; Bacescu, 1968 MED Caucasus and Crimea Coast, USSR Sheremetevskij, 1974 denticulatus ANE Fladden, UK Mclntyre, 1962 MED Naples, Italy Zelinka, 1928 diffussus MED Trieste, Italy Zelinka, 1928 * echinoderoides MED Naples, Italy Zelinka, 1928 ecphantor, ASW Twin Cays, Belize Higgins, this study new species egyptensis ISW Al-Ghardaqa, Egypt Higgins, 1966a emarginatus, ASW Twin Cays, Belize Higgins, this study new species flaveolalus ANE Oresund, Denmark Lang, 1936 ANE Gullmarfjord, Sweden Nyholm, 1947b; Nyholm and Nyholm, 1976 MED Trieste, Italy Zelinka, 1928 MED Naples, Italy Zelinka, 1928 NUMBER 18 79 TABLE 15.—Continued Species FAO sea area Locality Authority frequens ANW Mt. Desert IS. (Me.), USA Blake, 1930; Hyman, Higgins, 1965 1951; Buzzards Bay (Mass.), USA Wieser, 1960; Higgins , 1965 iniorhaptus, ASW Twin Cays, Belize Higgins, this study new species kielensis ANE Kiel, FRG Zelinka, 1928 ANE Oresund, Denmark Lang, 1936 longicornis, ASW Twin Cays, Belize Higgins, this study new species longihastatus MED Naples, Italy Zelinka, 1928 MED Trieste, Italy Zelinka, 1928 maximus ANE Kadelnnne, GDR Reimer, 1963 moderatus ANE W. Sweden Nyholm, 1947b MED Naples, Italy Zelinka, 1928 odhneri PSE Cumberland Bay, S. Georgia Is. Lang, 1949 ponticus MED Odessa, USSR Reinhard, 1881, 1885 1887 MED Naples, Italy Zelinka, 1928 MED Constanta, Rumania Bacescu and Bacescu, 1956 MED Tusaul, Rumania Bacescu and Bacescu, 1956 MED Chitic, Rumania Bacescu and Bacescu, 1956 MED Bulgarian coast Marinov, 1964 MED Danube Delta, Rumania Bacescu, 1968 MED Caucasus coast, USSR Sheremetevskij, 1974 rectilineatus ANE W. Sweden Nyholm, 1947b MED Naples, Italy Zelinka, 1928 robustus MED Naples, Italy Zelinka, 1928 MED Trieste, Italy Zelinka, 1928 rugosus MED Naples, Italy Zelinka, 1928 sanjuanensis INE Orcas Is. (Wash.), USA Higgins, 1961 sculptus PSE Berkeley Sound, Falkland Is. Lang, 1949 solidus MED Naples, Italy Zelinka, 1928 tenuis MED Naples, Italy Zelinka, 1928 validus MED Trieste, Italy Zelinka, 1928 zelinkaei ANE Clew Bay, Ireland Southern, 1914 ANE Fladden, UK Mclntyre, 1962 species ISW Andaman Is. Higgins and Rao, 1979 ponticus Marinov, 1964, with P. kielensis, separat- ing P. ponticus sensu Zelinka from P. ponticus Rein- hard, 1881, and then renaming P. ponticus sensu Zelinka P. neapolitanus. It is unfortunate that Reinhard's (1881, 1885, 1887) descriptions and particularly his illustra- tions (1885, figs. 19-42) and those of Bacescu (1968) and Marinov (1964) are of such limited use. Zelinka could only have been frustrated by Reinhard's illustrations and must have used the peculiar subcuticular scars of the penultimate sternal plates as an important criterion for con- sidering Reinhard's species conspecific with his own specimens, which were the basis for his de- scription and illustrations (Zelinka, 1928, pi. 11: fig. 11; pi. 13: figs. 5, 6). Bacescu's (1968, pi. 6) illustrations of P. neapolitanus are no more than a crude copy of Zelinka's excellent illustrations and, therefore, cannot be based on material seen by Bacescu. In his attempt to improve on Bacescu's descrip- tion and illustrations of P. neapolitanus, Shereme- teveskij (1974) produced illustrations (pi. 2: fig. 8-12) that are no better than those of Reinhard, and it seems inconceivable that the former au- thor's illustrations represent the same species con- sidered P. ponticus by Zelinka. In particular, there is no agreement in the number and position of sensory setae, nor do the shapes of either the entire trunk or the midventral plates bear any resemblance to one another. Until more convincing evidence emerges, I 80 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES FIGURE 191.—Distribution of Pycnophyes (solid circles indicate records of named species, stippled circle indicates genus records only). shall consider P. neapolitanus Bacescu, 1968, in- cluding the interpretation by Sheremetevskij (1974), a junior synonym of P. ponticus Zelinka, 1928. Pycnophyes flagellatus and P. quadndentatus have been synonymized herein and reassigned to Par- acentrophyes, leaving Pycnophyes echinoderoides as a species indeterminatum. It is very doubtful that any of the remaining 13 species of Pycnophyes, which have been described only from juvenile stages, can be matched with an adult. A compilation of the current status of the genus Pycnophyes is shown in Table 14. Table 15 and Figure 191 illustrate the reported distribution of these species. In number of described species, Pycnophyes with its total of 36, 24 of which are based on adults, is second only to Echinoderes with 65, 38 of which are based on adults. Because of both fewer records and number of species, little can be written about the distribution or ecology of this genus. Some, like Pycnophyes beaufortensis, have been found in estuaries (Higgins, 1964b) and the low salinities of the Black Sea and Baltic Sea (Sheremetevskij, 1974). Normally, members of this genus are associated with mud to sandy subtidal bottoms, rarely in algae. None have been found in association with other invertebrates. Pycnophyes represents the largest kinorhynchs; their maximum trunk length reaches 980 /xm in P. sculptus, and a minimum adult measurement is 380 /xm for P. ponticus. On this basis one would assume them to be easier to see and hence more commonly reported. In part, the apparent dis- parity in numbers of the homalorhagid genus Pycnophyes versus the cyclorhagid genus Echinoderes is an artifact of my own research procedure, for my own collections include many additional un- described species from the Arctic to the Antarctic, from intertidal estuarine mud, to high energy sandy beaches and abyssal sediments throughout the world. NUMBER 18 81 Genus Kinorhynchus Sheremetevskij, 1974 Key to Adults of Kinorhynchus 1. Middorsal processes present on segment 11 2 Middorsal processes absent on segment 11 8 2. Middorsal processes on segments 4-11, small, blunt or rounded; lateral protuberances of terminal segment bi- or trifurcate K. giganteus Zelinka, 1908 Middorsal processes spinose; lateral protuberances of terminal segment bulbose or pointed 3 3. Segment 12 with single lateral seta on each margin; middorsal processes of anterior trunk segments blunt, bell-shaped K. ilyocryptus Higgins, 1961 Segment 12 with 2 or 3 lateral setae on each margin; middorsal processes spinose 4 4. Segment 12 with 3 lateral setae on each margin K. trisetosus, new species Segment 12 with 2 lateral setae on each margin 5 5. First trunk segment with subdorsal setae K. anomalus (Lang, 1953) First trunk segment without subdorsal setae 6 6. Lateral setae absent on segments 5, 7, and 9 only; terminal segment slightly pointed; margin of terminal segment pointed K. spinosus (Lang, 1949) Lateral setae absent on other combination of segments; margin of terminal segment not pointed 7 7. Lateral setae absent on segments 3, 5, 6, 8, 9, and 11; segment 12 sharply tapered; terminal segment very narrow . K. stenopygus, new species Lateral setae absent on segments 5 and 11; segment 12 tapering only slightly; terminal segment broad . .. K. cataphractus (Higgins, 1961) 8. Lateral setae absent on segment 11 9 Lateral setae present on segment 11 13 9. Lateral setae present on segment 9 K. belizensis, new species Lateral setae absent on segment 9 10 10. Lateral setae absent on segment 10 K. paraneapolitanus Sheremetevskij, 1974 Lateral setae present on segment 10 11 11. Lateral setae absent on segment 6; segment 12 very short, bulging lateroterminally K. distentus, new species Lateral setae present on segment 6 12 12. Lateroterminal margins of segment 12 distinctly spinose, extending well beyond terminal segment; anterior margin of first trunk segment much narrower than that of segment 7; middorsal spinous process on segment 10 K. mainensis Blake, 1930 Lateroterminal margins of segment 12 spinose but not extending beyond terminal segment; anterior margin of first trunk segment about as wide 82 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES as that of segment 7; no middorsal process on segment 10 K. langi (Higgins 1964b) 13. Lateral setae missing on segment 12; diagonal buttresses extending from midlateral margins to pachycycli on segments 3-11 K. erismatus, new species Two lateral setae present on segment 11; no diagonal buttresses present • 14 14. Middorsal setae on segments 4-11; segment 12 much longer than segment 11; 2 lateral setae of segment 12 very prominent, near posterior margin K. apotomus, new species Middorsal setae absent; segment 12 much shorter than segment 11; 2 lateral setae of segment 12 small, not near posterior margin K. deirophorus, new species Kinorhynchus apotomus, new species FIGURES 192-203 DIAGNOSIS.—Trunk segments 3-9 nearly equal in width, segments 10-13 tapering moderately; anterodorsal margin of first trunk segment evenly dentate; prominent midorsal setae on segments 4-11; lateral setae on segments 4, 6-11, 2 on either side of segment 12 near terminal margin; terminal segment truncate; L-shaped cuticular scars posteriorly adjacent to sensory spots on ep- isternal plates. DESCRIPTION.—Adults (Figures 192-203), trunk length 592 /xm, segments 3-9 nearly equal in width, segments 10-13 tapering moderately; MSW-8 136-140 /xm, 23.0-23.6 percent of trunk length; SW 112 /xm, 18.6 percent of trunk length. Second segment with 4 dorsal and 4 ventral placids, margins even (Figures 194, 196). Trunk segments (Figures 192, 193) without middorsal spinose processes, subcuticular rounded middorsal spinose structure present on segments 4-11. Prominent setae, 22-24 /xm long, situated middorsally on segments 4-11, dorsolat- erally on segments 3-11, laterally on segments 4, 6-11 with 2 on either side of segment 12 near terminal margin, ventrolaterally on sternal plate of segments 3 and lateral plates, mesial to sensory spots, segments 5-10, more mesial on segment 11. Sensory spots subdorsal on first trunk segment, 2 subdorsal sensory spots on segments 4-12, sensory spot near anterior margin of each episternal plate of first trunk segment or lateral to setae or setal position on lateral plates of segments 4-12. Pa- chycycli weakly developed; midventral thicken- ings at anteromesial margins of segments 10-12; series of cuticular ridges beginning with longitu- dinal subdorsal ridges on tergal plate of segment 3 (Figure 196), continuing as transverse ridges, becoming more uneven, beginning at anterolat- eral margins of tergal and sternal plates 4-12 (Figures 192, 193). Prominent muscle scars dor- solateral and ventrolateral on first trunk segment, prominent L-shaped cuticular scar (Figure 194) below sensory spot on each episternal plate; cres- cent-shaped cuticular scar near midline of sternal plates, beginning with 45° orientation on anterior segments, becoming more transverse on posterior segments changing to longitudinal orientation on segment 12, tergal plates with crescent-shaped cuticular scars anterior to lateralmost sensory spot, scars nearly transverse in orientation on all segments. First trunk segment (segment 3) with antero- lateral margins of tergal plate extending to form slightly pointed projections, anterior tergal mar- gin evenly dentate (Figure 195), episternal plates with sculptured anteromesial area of thin cuticle, single such area at anterior margin of episternal plates (Figures 192, 194, 198); midsternal plate trapezoidal, about 86 /xm long, 70 /xm basal width tapering to a slightly expanded, rounded apex 26 /xm in width. Sternal plates of segment 4 of males with prom- inent adhesive tube, 35-37 /xm long (Figure 194). Pachycyclus extensive, with granular surface sculpturing. Segments 5-11 similar except for differences already noted. NUMBER 18 83 50um lOOum Segment 12 longer than segment 11, sternal plates incised near posterolateral margins (Figure 196), truncate at midline; tergal plate projecting slightly at midline. Terminal tergal segment broadly truncate, ex- tending slightly beyond penultimate segment, minutely pointed lateral terminal projections, about 8 /xm long, at margin (Figures 196, 197); terminal sternal margin appears parallel with preceding sternal margin; 1 pair of prominent penile spines and 1 or 2 pairs of smaller penile spines near anterolateral margins of terminal ster- nal plates (cuticular hairs occasionally create the impression that more penile spines are present, requiring this character to be treated with cau- tion). Morphometric data for the second of two males were identical for TL, SW, SW/TL. Only the MSW measurement differed (MSW 140 /xm, 23.6 percent of trunk length). HOLOTYPE.—Adult male, TL 592 /xm (Figures 192-203), Twin Cays, sta RH 444, Belize (16°50.0'N, 88°06.0'W), 8 Apr 1977, col. R.P. Higgins, USNM 70008. PARATYPE.—Adult male, TL 592 /xm, other data as for holotype, USNM 70009. REMARKS.—Kinorhynchus apotomus is unique in having middorsal setae. To be sure, setae are sometimes difficult to see, and previous descrip- tions could be faulty, but the middorsal setae on segments 4-11 are prominent as are all others on the trunk segments of this new species. With the exception of K. ilyocryptus, members of this genus have two lateral setae on segment 12. Kinorhynchus apotomus also has two, but they are located very close to the terminal margin, which emphasizes their prominence (Figures 195, 196). No previously described species of Kinorhynchus has posterolateral^ indented margins of the twelfth sternal plates (Figure 196) followed by the severely truncated terminal segment. The pointed FIGURES 192-197.—Kinorhynchus apotomus, new species: 192, neck and trunk segments, ventral view, holotypic male (USNM 70008); 193, same, dorsal view; 194, same, segments 2-4, lateral half, ventral view; 195, same, dorsal view; 196, same segments 12, 13, lateral half, ventral view; 197, same, dorsal view. 84 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES —> 10 198 ■ 201 ■ l I 199 I 202 ! 200 m \ 203 NUMBER 18 85 lateral terminal protuberances of the terminal segment are similar to those described for several species, including K. anomalus. These structures, more commonly bulbous or accompanied by one or more additional minute protuberances, have led a few authors to speculate as to the validity of the separation of Kinorhynchus from Pycnophyes. Blake (1930) predicted the ultimate union of these two genera, and Lang (1949) referred to these structures as lateral terminal spines but defended the separation of the genera. These protuberances so far do not appear to be articulated in Kinorhyn- chus as contrasted with those of Pycnophyes. The anterodorsal margin of K. apotomus is den- tate (Figure 196), a feature shared with K. gigan- teus. No other species has demonstrated the series of transverse parallel cuticular ridges that occur near the lateral margins of both the tergal and sternal plates of this new species. These ridges are similar to those of Pycnophyes corrugatus as noted earlier. The thin cuticular areas of the episternal and midsternal plates are similar in number and position to K. mainensis and K. ilyocryptus. The L- shaped cuticular scars posteriorly adjacent to the sensory spots on the episternal plates have not been observed in other species. ETYMOLOGY.—The name of this species is from the Greek apotomus (cut off, abrupt). Kinorhynchus belizensis, new species FIGURES 204-219 DIAGNOSIS.—First trunk segment slightly nar- rower than second trunk segment, maximum width at segment 7, tapering moderately there- after, terminal segment slightly indented at mid- line; midsternal plate of first trunk segment with nearly parallel lateral margins; middorsal setae at least on segment 8-12; reduced middorsal spinose protuberances on segments 3, 5-9; no lateral setae on segment 12; buttress-like cuticular FIGURES 198-203.—Kinorhynchus apotomus, new species: 198, segments 2-4, ventral view; 199, segments 7, 8, ventral view; 200, segments 11-13, ventral view; 201, segments 2-4, dorsal view; 202, segments 7, 8, dorsal view; 203, segments 12, 13, dorsal view. (Interference contrast photographs all with same scale (in /tm) as shown in Figure 198.) support present between lateral margin and pa- chycyclus of each sternal plate. DESCRIPTION.—Adults (Figures 204-219), trunk length, 478-614 /xm, first trunk segment slightly narrower than second trunk segment, MSW-7 136-168 /xm, 22.2-33.5 percent of trunk length; trunk moderately tapered, SW 92-136 /xm, 18.3-27.6 percent of trunk length. Second trunk segment with 4 dorsal and 2 ventral placids, dorsomesial placid with slightly indented margin (Figures 206, 207). Trunk segments (Figures 204, 205) with small, 22-23 /xm, middorsal spinose processes at poste- rior margins of segments 3, 5-9, slightly inset into margin on segments 3, 5-7. Setae, 20-21 /xm long, situated middorsally on at least segments 8-12 (several specimens, including holotype, show off- centered setae on segments 4, 6, and 7; these may not occur in all specimens, see Figure 205); sub- dorsal setae on segments 7-11, laterodorsal setae on segments 10, 11, lateral setae on segments 3- 10, none on 11 or 12, ventrolateral setae on segments 6-11; sensory spots subdorsal on seg- ments 4-11, dorsolateral on segments 3, 4, 6-8, 10; additional sensory spots lateral to dorsolateral on segment 8; 3 sensory spots on each episternal plate; ventral sensory spots on each sternal plate 4-7; 2 ventral spots on sternal plates 8-11; single ventral spot on each sternal plate of segment 12. Pachycycli well developed, especially those of segment 11 (Figures 216, 219) where midventral thickenings appear to be laterally displaced. Slightly ridged, longitudinally oriented pattern evident on episternal plates (Figures 206, 210), 2 transverse, parallel ridges near pachycycli of ster- nal plates 4-5, 1 ridge diverting as a posterior loop evident on sternal plates 5-11 (Figure 204). Muscle scars lateral on tergal plate of segment 3; scars continuing across anterior third of plate in pattern shown in Figure 205; cuticular scars somewhat triangular in shape, lateral to subdorsal sensory spots on tergal plates 4-11, becoming more anteriorly situated in posterior segments, anterolateral to middorsal setae on segment 12, lateral to ventral midline on sternal plates, pos- teriorly adjacent to anterormost sensory spots of episternal plates. 86 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES lOOjjm 50jum 212 213 FIGURES 204-213.—Kinorhynchus belizensis, new species: 204, neck and trunk segments, ventral view, holotypic male (USNM 70010); 205, same, dorsal view; 206, same, segments 2-4, lateral half, ventral view; 207, same, dorsal view; 208, same, segments 12, 13, lateral half, ventral view; 209, same, dorsal view; 210, segments 2-4, lateral half, ventral view, allotypic female (USNM 70011); 211, same, dorsal view; 212, same, segments 12, 13, lateral half, ventral view; 213, same, dorsal view. FIGURES 214-219.—Kinorhynchus belizensis, new species: 214, segments 3, 4, ventral view; 215, segments 7, 8, ventral view; 216, segments 11-13, ventral view; 217, segments 3, 4, dorsal view; 218, segments 7, 8, dorsal view; 219, segments 11-13, dorsal view. (Interference contrast photographs all with same scale (in /im) as shown in Figure 214; BU = buttress.) SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES TABLE 16.—Measurements (/xm) and indices (%) for Kinorhynchus belizensis adults (see "Methods" for character abbreviations) Charact* :r Number Range Mean Standard deviation Standard error Coefficient of variability TL 6 8 478-592 543.0 42.4 15.0 7.8 2 12 478-614 555.3 42.8 12.4 7.7 62 20 478-614 550.4 42.0 9.4 7.6 SW 6 8 112-124 119.5 4.0 1.4 3.3 2 12 92-136 125.3 11.4 3.3 9.1 62 20 92-136 123.0 9.4 2.1 7.7 SW/TL 6 8 18.3-24.9 22.4 2.3 0.8 10.11 2 12 19.2-27.6 22.6 2.2 0.6 9.7 62 20 18.3-27.6 22.5 2.2 0.5 9.6 MSW-7 6 8 136-156 150.5 6.4 2.3 4.3 2 12 148-168 162.7 6.2 1.8 3.8 62 20 136-168 157.8 8.7 1.9 5.5 MSW/TL 6 8 22.2-31.8 27.9 2.8 1.0 10.1 2 12 25.9-33.5 29.4 1.9 0.6 6.5 62 20 22.2-33.5 28.8 2.4 0.5 8.3 First trunk segment (segment 3) with only slight projection of the anterolateral tergal margins; anterior tergal margin roughly denticulate (Fig- ures 207, 211). Midsternal plate long, 96 /xm, narrow, 52 /xm, lateral margins nearly parallel; episternal plates with apparent partial longitu- dinal division created by inner area of thicker cuticle. First trunk segment slightly shorter in female than in male (Figures 210, 211). Sternal plates of segment 4 of males with ad- hesive tubes, 38 /xm long (Figure 206); females lack adhesive tubes, have smaller sensory spots, and the elongated cuticular scars are in a more posterior position; ventral accessory pachycycli narrow, arching at primary pachycycli. Middor- sal protuberance not evident on posterior margin. Segment 5 without ventral setae, occasionally with 1 or more subdorsal setae. Segment 6 with ventral setae and sensory spots as noted earlier, segments 7-10 similar, trunk begins to taper be- ginning with segment 7 or 8. Segment 11 with enlarged pachycycli and laterally displaced mid- ventral thickenings. Segment 12 narrow at pos- terior margin, without lateral setae, only a serrate lateral margin partially caused by the distal ends of cuticular scales that appear like small hairs in lateral view. Terminal tergal segment indented at midline, giving a slightly bilobed appearance to the mar- gin, small bulbous projection at lateral margins, margins of terminal sternal plates parallel those of segment 12 (Figures 208, 209, 212, 213), males with 3 pairs of penile spines at anterolateral margins of terminal sternal plates (Figure 204), females with single gonopore (Figure 212). Males differ from females by presence of 3 pairs of penile spines at the anterolateral margins of terminal sternal plates; adhesive tubes on sternal plates of segment 4; slightly longer first trunk segment; variable presence of middorsal and sub- dorsal setae, especially in the anterior region of the trunk; slightly narrower trunk. Morphometric data for adult specimens are shown in Table 16. HOLOTYPE.—Adult male, TL 592 /xm (Figures 204-209, 214-219), Twin Cays, sta RH 442, Be- lize (16°50.0'N, 88°06.0'W), 8 Apr 1977, col. R.P. Higgins USNM 70010. ALLOTYPE.—Adult female, TL 572 /xm (Figures 210-213), other data as for holotype, USNM 70011. PARATYPES.—Six adult males and 10 females, TL 478-614 /xm, other data as for holotype, USNM 70012; 1 male, TL 614 /xm, Twin Cays, NUMBER 18 89 sta RH 443, other data as for holotype, USNM 70013; 1 female, TL 478 /xm, Twin Cays, sta RH 444, other data as for holotype, USNM 70014. DISCUSSION.—The most noticeable characters of Kinorhynchus belizensis are the slightly inset mid- dorsal spinose projections at the posterior margins of segments 3, 5-7 (Figure 205), the prominent pachycycli of segment 11 (the latter character is coupled with distinct midventral thickenings which are laterally displaced from their normal position, Figures 216, 219), and the buttress-like reinforcements of the pachycycli of the sternal plates (Figure 204). Since all previously described species of Kinorhynchus have two, or less commonly one, lateral setae on segment 12, the absence of any such setae in K. belizensis is another character of note. In general trunk shape, K. belizensis is similar to K. langi; the latter species also has poorly devel- oped middorsal spinous projections, restricted to the first five or six trunk segments as in K. belizen- sis, and both have four dorsal and only two ventral placids. In this latter point, however, the presence of two or four ventral placids is a matter of some subjectivity, since the thickness of the cuticle may vary only slightly, and in cases where the lateral placid area is not as thick as the mesial placids, one may judge only the two thicker placids as the only ones present. ETYMOLOGY.—This species is named after the country where it was discovered. Kinorhynchus deirophorus, new species FIGURES 220-235 DIAGNOSIS.—First trunk segment almost of equal width, maximum width at segment 7, ta- pering moderately; anterodorsal margin even with row of punctate sculpturing followed by an undulating ridge of cuticle, anterior margin of episternal plates with 2 areas of sculptured, thin cuticle, lateral area slightly anterior to mesial area, 2 similar areas at margin of midsternal plates, middorsal ridge of cuticle prominent on anterior trunk segments; prominent ventral pa- chycycli and laterally displaced midventral thick- enings on segment 11, segment 12 one-half as long as segment 11. DESCRIPTION.—Adults (Figures 220-235), trunk length, 592 /xm, segment 3 only slightly narrower at anterior margin than next few segments, MSW-7 128-180 /xm, 23.3-28.5 percent of trunk length; trunk tapering moderately, SW 116-136 /xm, 16.7-21.2 percent of trunk length. Second segment with 4 dorsal and 2 ventral placids, margins even (Figures 222, 223, 226, 227). Trunk segments (Figures 220, 221) without middorsal projections. Slight cuticular ridge on segments 3-11 and slight indication of spinose subcuticular structure on segments 3-9. Setae, 20-23 /xm long, situated dorsolaterally on seg- ments 5-11, laterally on episternal plate, laterally on sternal plates 4-11, with 2 setae on either side of segment 12, ventral setae absent. Sensory spots subdorsal on segments 3, 6, 8, 10, 11, dorsolateral on segments 5-12 (mesial to setae), ventral on episternal plates and sternal plates 4-12. Pachy- cycli well developed, especially on segments 4 and 11 (Figures 230, 232, 233, 235); midventral thick- enings of segment 11 laterally displaced (Figure 232); muscle scars with distinctive pattern on tergal plate (Figures 223, 227) and episternal plates (Figures 222, 226) of first trunk segment. Small, round to oval cuticular scars on either side of midline on sternal and tergal plates 4-11, (Figures 220, 221). Sternal plates 4-11 with single ridge of cuticle parallel with pachycycli laterally, extending posteriorly, a wide arc mesially, espe- cially on anterior segments. First trunk segment (segment 3) with distinct, somewhat blunt projections of the anterolateral tergal margins; anterior tergal margin even, with row of punctate sculpturing followed by an un- dulating ridge of cuticle (Figures 223, 227). Mid- dorsal ridge of cuticle on posterior half of tergal plate; anterior margins of episternal plates with 2 areas of sculptured, thin cuticle, lateral area slightly anterior to mesial area, 2 similar areas at margin of midsternal plate; midsternal plate trap- ezoidal, lateral margins undulant, length about 100 /xm, basal width 82 /xm, 48 /xm at apex. Sternal plates of segment 4 of males each with adhesive tube, 34-36 /xm long (Figure 227); fe- 90 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES Jum 227 IS FIGURES 220-229.—Kinorhynchus deirophorus, new species: 220, neck and trunk segments, ventral view, holotypic male (USNM 70015); 221, same, dorsal view; 222, same, segments 2-4, lateral half, ventral view; 223, same, dorsal view; 224, same, segments 12, 13, lateral half, ventral view; 225, same, dorsal view; 226, segments 2-4, lateral half, ventral view, allotypic female (USNM 70016); 227, same, dorsal view; 228, same, segments 12, 13, lateral half, ventral view; 229, same, dorsal view. FIGURES 230-235.—Kinorhynchus deirophorus, new species: 230, segments 3, 4, ventral view; 231, segments 7, 8, ventral view; 232, segments 11-13, ventral view; 233, segments 3, 4, dorsal view; 234, segments 7, 8, dorsal view; 235, segments 11-13, dorsal view. (Interference contrast photographs all with same scale (in jum) as shown in Figure 230.) 92 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES TABLE 17.—Measurements (/xm) and indices (%) for Kinorhynchus deirophorus adults (see "Methods" for character abbreviations) Character Number Range Mean Standard deviation Standard error Coefficient of variability TL 6 10 655-728 689.4 22.9 7.3 3.3 2 12 603-717 656.8 37.3 10.8 5.7 62 22 603-728 671.6 35.1 7.5 5.2 SW 6 10 120-132 126.0 4.7 1.5 3.7 2 12 116-136 128.0 6.2 1.8 4.8 62 22 116-136 127.1 5.5 1.2 4.3 SW/TL 6 10 16.7-19.5 18.3 0.9 0.3 4.9 2 12 18.4-21.2 19.6 0.9 0.3 4.8 62 22 16.7-21.2 19.0 1.2 0.3 6.2 MSW-7 6 10 160-176 167.6 5.2 1.6 3.1 2 12 128-180 166.3 14.2 4.1 8.6 62 22 128-180 166.9 10.8 2.3 6.5 MSW/TL 6 10 23.3-25.4 24.3 0.6 0.2 2.6 2 12 21.2-28.5 25.3 2.0 0.6 7.7 62 22 21.2-28.5 24.9 1.6 0.3 6.3 males lack adhesive tubes but have setae in the same location. Segments 5-11 similar except for characters already noted. Segment 12 noticeably smaller than segment 11, about one-half the length of the latter; pos- terior margin of sternal plates nearly even, pos- terior margin of tergal plate slightly pointed at midline in male, less so in female (Figures 225, 229). Terminal segment small, sternal margins even, tergal margin with lateral spinous projection in male (Figure 224), 1 or 2 additional marginal interruptions, more mesial in the female (Figure 228), broadly rounded in both sexes. One large and 2 small penile spines at anterolateral margin of each sternal plate (Figure 284); single gonopore in females (Figure 228). Males differ from females by presence of 3 pairs of penile spines at anterolateral margins of ter- minal sternal plates, adhesive tubes on sternal plates of segment 4, and slightly more pro- nounced sculpturing on first trunk segment (Fig- ures 223, 227). Morphometric data for adult specimens are shown in Table 17. HOLOTYPE.—Adult males, TL 697 /xm (Figures 220-225, 230-234), Twin Cays, sta RH 442, Be- lize (16°50.0/N, 88°06.0'W), 8 Apr 1977, col. R.P. Higgins, USNM 70015. ALLOTYPE.—Adult female, TL 707 /xm (Figures 226-229), other data as for holotype, USNM 70016. PARATYPES.—Nine males and 11 females, TL 603-728 /xm, other data as for holotype, USNM 70017. DISCUSSION.—The pattern of cuticular sculp- turing on the anterior margin of tergal plate 3 is the most obvious of the characters which separate K. deirophorus from all previously described mem- bers of this genus. This character resembles the tergal plate sculpturing of Pycnophyes iniorhaptus from this same locality. Kinorhynchus paraneapoli- tanus and K. apotomus, like K. deirophorus, lack middorsal processes at the posterior margin of at least some trunk segments. A second unique character is the pattern of two thin, sculptured areas at the anterior margin of both the two episternal and single midsternal plates of the first trunk segment. The only other known species with two such areas on the epister- NUMBER 18 93 nal plates is K. giganteus. This latter species ap- pears to have three less well-defined areas on the midsternal plate. The prominent pachycycli of segment 11 are similar to those previously de- scribed for K. belizensis; also similar are the lat- erally displaced midventral thickenings. Like the latter species, K. deirophorus has a ridge pattern on the sternal plates which suggests a further simi- larity. ETYMOLOGY.—The name of this species is from the Greek deiropedes (necklace) plus phoras (bear- ing), in reference to the sculptured margin of the tergal plate of the first trunk segment. Kinorhynchus distentus, new species FIGURES 237-257 DIAGNOSIS.—Trunk only slightly narrowed both anteriorly and posteriorly; tergal plate of first trunk segment with 5 looped ridges on either side of midline; 2 thin cuticular areas at margin of episternal and midsternal plates; lateral mar- gins of segment 11 extended posteriorly; penulti- mate segment short, one-half the length of the preceding segment, expanded and distended lat- erally, forming rounded margin; terminal seg- ment with spinose lateral extensions of the mar- gins; distinctive cuticular fringe pattern near mid- line of sternal plates on segment 4; series of 1 to 3 parallel cuticular ridges extending mesially from lateral margins of sternal plates 4-12. DESCRIPTION.—Adults (Figures 236-257), trunk length 540-665 /xm, trunk only slightly narrowed both anteriorly and posteriorly; MSW-6 152-168 /xm, 24.7-29.6 percent of trunk length; SW 112- 120 /xm, 18.0-20.7 percent of trunk length. Sec- ond segment with 4 dorsal and 2 ventral placids, margins even (Figures 236, 242, 246, 252). Trunk segments (Figures 236, 237) without middorsal projections but with slight evidence of subcuticular spinose structure on segments 3-9. Setae, 20-22 /xm long, situated dorsolaterally on segments 3-11, laterally on episternal plates and sternal plates 4, 7, 10, and 2 on lateral margins of segment 12; ventral setae near lateral margins of episternal plates and each sternal plate of seg- ments 5, 8-10, those of segment 5 lateral to sensory spot, others mesial to sensory spot (Figure 236). Sensory spots subdorsal on segments 3, 4, 6, 8, 10, 11; 1 to 3 dorsolateral sensory spots on other segments; ventral sensory spot near anterior margin of episternal plate and on sternal plates 4-12. Series of 1 to 3 parallel cuticular ridges extending mesially from lateral margins of sternal plates 4-12. Pachycycli well developed, particu- larly on segments 4, 10, and 11, midventral thick- enings on sternal plates of segments 10 and 11 (Figures 236, 248). Large muscle scars dorsolat- erally on tergal plate and on episternal plates of segment 3; crescent-shaped cuticular scars, lon- gitudinally oriented, posterior to sensory spot, mesial to setae of episternal plates; small, round to oval cuticular scars on sternal plates 4-12 near ventral midline, small, round to triangular- shaped cuticular scars on segments 4-12 near lateral margins of tergal plate 4, nearer to dorsal midline on tergal plates 5-12. First trunk segment (segment 3) with moderate projections of the anterolateral tergal margins, anterior tergal margin with slightly roughened sculpture followed by series of 5 looped ridges on either side of midline (Figure 237). Anterior mar- gin of episternal plates with 2 areas of sculptured, thin cuticle, lateral area slightly anterior to mesial area, 2 similar areas at margin of midsternal plate (Figures 236, 238, 252); midsternal plate trape- zoidal, about 87 /xm long, 68 /xm basal width, 38 /xm wide at apex. Sternal plates of segment 4 of males with short, 25-30 /xm, adhesive tubes (Figure 238); females lack adhesive tubes, and some lack dorsolateral setae on this segment (Figure 243). Two parallel ridges of fringe (Figures 238, 254) present, ex- tending below adhesive tubes from lateral mar- gins to midventral line. Segments 5-10 similar except for characters already noted. Segment 11 with distinctly projecting lateral margins (Figures 248, 257), margin fringed (Figure 256) but only visible through use of scanning electron micro- scope. 94 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 50/JID FIGURES 236-245.—Kinorhynchus distentus, new species: 236, neck and trunk segments, ventral view, holotypic male (USNM 70018); 237, same, dorsal view; 238, same, segments 2-4, lateral half, ventral view; 239, same, dorsal view; 240, same, segments 12, 13, lateral half, ventral view; 241, same, dorsal view; 242, segments 2-4, lateral half, ventral view, allotypic female (USNM 70019); 243, same, dorsal view; 244, same, segments 12, 13, lateral half, ventral view; 245, same, dorsal view. HI T * 249 I s- #" ^ 1 i >£fc i 247 ^ s ( 1 i \ m 250 248 251 FIGURES 246-251.—Kinorhynchus distentus, new species: 246, segments 3, 4, ventral view, holotypic male (USNM 70018); 247, same, segments 7, 8, ventral view; 248, same, segments 11-13, ventral view; 249, same, segments 3, 4, dorsal view; 250, same, segments 7, 8, dorsal view; 251, same, segments 11-13, dorsal view. (Interference contrast photographs all with same scale (in /im) as shown in Figure 246.) 96 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES NUMBER 18 97 TABLE 18.—Measurements (jum) and indices (%) for Kinorhynchus distentus adults (see "Methods" for character abbreviations) Character" Standard Standard Coefficient Number Range Mean deviation error of variability TL 6 1 614 0.0 0.0 0.0 0.0 2 3 540-665 606.3 62.9 36.3 10.4 62 1 540-665 608.3 51.5 25.7 8.5 SW 6 1 112 0.0 0.0 ().() 0.0 2 3 112-120 114.7 4.6 2.7 4.0 62 4 112-120 114.0 4.0 2.0 3.5 SW/TL 6 1 18.2 0.0 0.0 ().() 0.0 2 3 18.0-20.7 19.0 1.5 0.9 7.8 62 4 18.0-20.7 18.8 1.3 0.6 6.8 MSW-6 6 1 152 0.0 0.0 0.0 0.0 2 3 160-168 164.0 4.0 2.3 2.4 62 4 152-168 161.0 6.8 3.4 4.2 MSW/TL 6 1 24.7 0.0 0.0 0.0 0.0 2 3 25.3-29.6 27.2 2.2 1.3 8.0 62 4 24.7-29.6 26.6 2.2 1.1 8.2 Segment 12 short, one-half the length of the preceding segment, expanded and distended lat- erally forming rounded margins. Midventral area of sternal plates projects posteriorly. Terminal tergal plate with distinct, thin projection of lateral margin (Figures 240, 244, 257); sternal plates with 2, possibly 3, pairs of penile spines near anterolateral margins in males, females with sin- gle gonopore. Males differ from females by presence of 2 (or 3) pairs of penile spines at the anterolateral mar- gins of terminal sternal plates, presence of adhe- sive tubes on sternal plates of segment 4, and some minor variation in dorsolateral setae pres- ence or position. Only one male and three females were avail- able for type material (one specimen used for SEM study); morphometric data (Table 18) may not be meaningful. FIGURES 252-257.—Kinorhynchus distentus, new species: 252, segments 2, 3, ventral view, male; 253, ventral sensory spot (enlarged from Figure 254), segment 4, left sternal plate, male; 254, segment 4, ventral view, left sternal plate, male; 255, segment 7, ventral view, right sternal plate, male; 256, segment 7, ventral view, portion of left sternal plates, male; 257, segments 11, 12, 13, ventral view, left sternal plates, male. (SEM photographs, each to scale (in /tm) indicated.) HOLOTYPE.—Adult male, TL 614 /xm, (Figures 236-241, 246-251), Twin Cays, sta RH 444, Be- lize (16°50.0'N, 88°06.0'W), 8 Apr 1977, col. R.P. Higgins, USNM 70018. ALLOTYPE.—Adult female, TL 665 /xm (Figures 242-245), other data as for holotype, USNM 70019. PARATYPES.—Two females, TL 540-614 /xm, other data as for holotype, USNM 70020. DISCUSSION.—Kinorhynchus distentus resembles K. deirophorus in the character of its thin sculptured areas on the anterior margin of the episternal and midsternal plates and K. apotomus in that both have a similar series of parallel cuticular ridges which are especially distinct near the lateral mar- gins of the anterior sternal plates. Furthermore, K distentus shares the otherwise unique short pen- ultinate segment character previously described for K. deirophorus; both of these latter species have marginal sculpturing on the tergal plate of the first trunk segment. This sculpturing is different in the two species but occurs in the same two areas and suggests a kinship. This species has very few characters which can be meaningfully com- pared with other species in the same genus. ETYMOLOGY.—The name of this species is from the Greek distentus (swell out, distended), which 98 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES reflects the character of the penultimate trunk segment. Kinorhynchus erismatus, new species FIGURES 258-273 DIAGNOSIS.—Trunk slightly tapered anteriorly, moderately so with rounded margin posteriorly; tergal plate of first trunk segment with minute projections of anterolateral margins, slightly den- ticulate at anterior margin; area of thin, sculp- tured cuticle across anterior margin of episternal plates, 2 such areas at anterior margin of mid- sternal plate; buttress-like reinforcement between lateral pachycycli of sternal plates 3-11; 1 to 3 parallel cuticular ridges along anterior half of sternal plates 4-11; 12th segment without lateral setae; terminal tergal plate with lateral bulbous projections. DESCRIPTION.—Adults (Figures 258-273), trunk length 614-770 /xm, trunk slightly tapered ante- riorly-, moderately so with rounded margin pos- teriorly; MSW-7 168-220 /xm, 22.8-30.9 percent of trunk length; SW 140-180 /xm, 18.7-27.4 per- cent of trunk length. Second segment with 4 dorsal placids with slightly dentate margins and 2 even margined ventral placids (Figures 260, 261, 264, 265). Trunk segments (Figures 258, 259) without middorsal marginal projections; slight evidence of subcuticular spinose structure on trunk seg- ments 4-11. Setae, 20-22 /xm long, situated sub- dorsally on segments 4-9, 12, dorsolaterally on segments 4-11, at extreme lateral margins of segments 3, 4 (but not always visible), laterally on episternal plates and sternal plates of segments 9 and 11, ventrally on sternal plates 6-11; setae lateral to ventral sensory spot on segment 6, mesial on segments 7-11; 1 to 3 parallel cuticular ridges along anterior half of sternal plates 4-11. Subdorsal sensory spots on segments 3-11; dor- solateral sensory spots on segments 4-12 with extra, more lateral, sensory spot on segment 10 (Figure 259); 2 additional sensory spots on the 12th tergal plate of female (Figure 267), 2 ventral sensory spots on episternal plates, 1 anterior, 1 posterior (Figure 264), single lateral sensory spot on each sternal plate of segments 4-12. Large muscle scars dorsolateral on tergal plate of first trunk segment, more visible on episternal plates; small, longitudinal cuticular scar posterior to an- teriormost sensory spots of episternal plates; small, oval to triangular cuticular scars usually anteriorly adjacent to dorsolateral setae of tergal plates 4-12; small, oval cuticular scars posteriorly adjacent to cuticular ridge on sternal plates 3-11, slightly larger, slightly crescent-shaped cuticular scar anterior to ventral sensory spots of penulti- mate sternal plates. Pachycycli moderately devel- oped, midventral thickenings absent. Buttress-like reinforcement between lateral margin and pachy- cycli of sternal plates 3-11. First trunk segment (segment 3) with minute projections of the anterolateral tergal margins, anterior tergal margin slightly denticulate. Mus- cle scars as illustrated (Figure 261). Area of thin sculptured cuticle across anterior margins of ep- isternal plates, 2 such areas at anterior margin of midsternal plate; midsternal plate trapezoidal, lateral margins slightly curved inward, length about 104 /xm, basal width 80 /xm, width at apex, 48 /xm. Sternal plates of segment 4 of males with ad- hesive tubes, about 43 /xm long (Figure 260); females lack adhesive tubes but have setae in same area (Figure 264). Accessory pachycycli complex (Figure 260); cuticular ridge looping posteriorly around ventral sensory spot of each sternal plate; with exception of sternal plates of segment 4, segments 4-11 are similar except for details already noted. Segment 12 tapering sharply, sternal margins bowed (Figure 262) occasionally obscuring ter- minal segment (Figure 266). Lateral setae lacking, replaced by 2 minute protuberances (possibly secretions) on either side of segment (Figure 262, 266). Terminal segment with bulbous projection at lateroterminal margin of tergal plate. Sternal plates less rounded, with 3 pairs of penile spines at anterolateral margins in males (Figures 262, 270), single gonopore in females. NUMBER 18 99 lOOLim I « 8 f • ' • T • " \ T "f •' o To 1 50pm FIGURES 258-267.—Kinorhynchus erismatus, new species: 258, neck and trunk segments, ventral view; holotypic male (USNM 70021); 259, same, dorsal view; 260, same, segments 2-4, lateral half, ventral view; 261, same, dorsal view; 262, same, segments 12, 13, lateral half, ventral view; 263, same, dorsal view; 264, segments 2-4, lateral half, ventral view, allotypic female (USNM 70022); 265, same, dorsal view; 266, same, segments 12, 13, lateral half, ventral view; 267, same, dorsal view. 100 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES NUMBER 18 101 TABLE 19.—Measurements (/xm) and indices (%) for Kinorhynchus erismatus adults (see "Methods" for character abbreviations) Standard Standard Coefficient Characte Number Range Mean deviation error of variability TL 6 25 624-770 718.8 37.9 7.6 5.3 2 19 614-760 698.6 40.6 9.3 5.8 62 44 614-770 710.4 39.8 6.0 5.6 SW 6 25 140-172 159.4 6.7 1.3 4.2 2 19 164-180 170.3 6.3 1.5 3.7 62 44 140-180 164.1 8.5 1.3 5.2 SW/TL 6 25 18.7-26.2 22.2 1.6 0.3 7.1 2 19 22.0-27.4 24.5 1.7 0.4 6.8 62 44 18.7-27.4 23.2 2.0 0.3 8.4 MSW-7 6 25 168-220 190.0 11.0 2.2 5.8 2 19 172-204 195.3 7.2 1.7 3.7 62 44 168-220 192.1 9.8 1.5 5.1 MSW/TL 6 25 22.8-30.2 26.4 1.7 0.3 6.5 2 19 23.6-30.9 28.0 1.7 0.4 6.1 62 44 22.8-30.9 27.1 1.9 0.3 6.9 Males differ from females by presence of 3 pairs of penile spines at anterolateral margins of ter- minal sternal plates, single sensory spot on the tergal plate of segment 12, presence of adhesive tubes on sternal plates of segment 4, and slightly narrower sternal width (Table 19). Morphometric data for adults of Kinorhynchus erismatus, new species, are shown in Table 19. HOLOTYPE.—Adult male, TL 749 /xm (Figures 258-263, 268-273), Twin Cays, sta RH 444, Be- lize (16°50.0'N, 88°06.0'W), 8 Apr 1977, col. R.P. Higgins, USNM 70021. ALLOTYPE.—Adult female TL 728 /xm (Figures 244-267), other data as for holotype, USNM 70022. PARATYPES.—Eleven males and 8 females, TL 655-770 /xm, Twin Cays, sta RH 443, other data as for holotype, USNM 70024; 10 males and 10 females, TL 614-749 /xm, Twin Cays, sta RH FIGURES 268-273.—Kinorhynchus erismatus, new species: 268, segments 2-4, ventral view; 269, segments 7, 8, ventral view; 270, segments 11-13, ventral view; 271, segments 2-4, dorsal view; 272, segments 7, 8, dorsal view; 273, segments 11-13, dorsal view. (Interference contrast photographs all with same scale (in jttm) as shown in Figure 268; BU = buttress.) 444, other data as fc; holotype, USNM 70025; 3 males, TL 697-730 /xm, Twin Cays, sta RH 442, other data as for holotype, USNM 70023., DISCUSSION.—Kinorhynchus ensimatus most closely resembles K. belizensis on the basis of the buttress-like reinforcement of the pachycycli, a character unique to these two species. Another similarity exists in that, likewise, these two species uniquely lack lateral setae on the penultimate segment. Both have a similar trunk shape, both have a similar sternal cuticular sculpturing, and both exhibit similar muscle scars on the tergal plate of the first trunk segment. There are differ- ences in accessory pachycycli structure between the two species, differences in the pachycycli of the 11th segment, as well as differences in setae and sensory spot arrangement, but these latter characters do not diminish the suggested close kinship. Although it is not similar in the same sense as K. belizensis, K langi partially resembles K. eris- matus in general shape and accessory pachycycli structure. Like most of the new species of this genus being described in this paper, there are many unique taxonomic characters and only a few general similarities between the Belizian 102 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES fauna and that of the remaining world. ETYMOLOGY.—This species name is from the Greek erismatus (buttressed), which reflects the structural reinforcement of the sternal pachycycli. Kinorhynchus stenopygus, new species FIGURES 274-301 DIAGNOSIS.—Trunk slightly tapered anteriorly, strongly tapered posteriorly with narrow penul- timate and slightly flaring terminal segments; well-developed middorsal spinose projections; sternal plates with prominent semicircular cuti- cular ridge; projections of the anterolateral tergal margins of the first trunk segment distinctly rounded; anterior tergal margin with 3 minute projections. DESCRIPTION.—Adults (Figures 274-301), trunk length 509-603 /xm, trunk slightly tapered ante- riorly, strongly tapered posteriorly; MSW-8 132- 152 /xm, 24.5-28.8 percent of trunk length; SW 96-112 /xm, 16.6-22.0 percent of trunk length. Second segment with 2 even-margined ventral placids, 4 dorsal placids suspected but not con- firmed (Figures 276-280). Trunk segments (Figures 274, 275) with spinose middorsal projections on posterior margins of segments 3-11. Setae, 19-22 /xm long, situated middorsally (or slightly off center) on segments 6 and 8 (occasionally on other segments), occasion- ally subdorsally on segment 3 (Figure 298), dor- solaterally on segments 3, 2 on 4, 5-11, lateral on episternal plate of segment 3, lateral on segments 7, 10, and 2 on segment 12, ventrally, centered on sternal plates of segments 4-11. Several small cuticular ridges present near anterolateral mar- gins of sternal plates on segment 4; prominent semicircular cuticular ridge centered on sternal plates of segments 3-11 (Figures 274, 292). Perispinal sensory spots on either side of middor- sal marginal projections of segments 3-11 (Fig- ures 274, 297, 299); 2 subdorsal sensory spots on segments 3-11, dorsolateral sensory spots near margin of segments 3, 5-11; ventral sensory spots anterolateral on episternal plates of segment 3 and near center of each sternal plate of segments 4-12, usually laterally adjacent to ventral setae but occasionally reversed, especially on segments 5 and 7. Prominent oval muscle scars on epister- nal plates (Figures 276, 280) and lateral on tergal plate of segment 3 (Figures 277, 281); elongate transverse cuticular scars dorsolateral on seg- ments 4-11, same scars more triangular-shaped on segment 12; oval to crescent-shaped cuticular scars posterior to semicircular cuticular ridges of sternal plates 4-11, in same relative position (ridges absent) on segment 12. Pachycycli well developed, narrow midventral thickenings on seg- ment 11. Surface of trunk segments with prominent scales on tergal plates (Figure 301) and lateral surfaces of most sternal plates, scales becoming more hairlike mesially and on posterior segments (Figure 295) but clearly visible only when using a scanning electron microscope. First trunk seg- ment (segment 3) with rounded projections of the anterolateral tergal margins, tergal margin even except for middorsal and lateral slightly protrud- ing points (Figures 277, 281, 298). Muscle scars, setae, and sensory spots as previously noted. No areas of thin cuticle present on episternal or mid- sternal plates; midsternal plate trapezoidal, lat- eral margins straight, length about 78 /xm, basal width 67 /xm, 40 /xm at apex, slight transverse line present one-fourth distance posterior. Sternal plates of segment 4 of males with ad- hesive tubes 37 /xm long (Figures 276, 291, 292); females lack adhesive tubes; several small cuti- cular ridges present at anterolateral margins, with prominent semicircular cuticular ridge poste- riorly adjacent to adhesive tubes (Figures 291, 292). Segments 5-11 similar with exceptions as pre- viously noted. Segment 12 strongly tapered, with rounded posterior margin. Lateral setae promi- nent in male, modified in female (Figures 278, 282,294,295). Terminal segment of both sexes small; female with lateral projection of tergal plate (Figures 282, 295), none in male (Figures 278, 294). Sternal plates small, often hidden; tergal plate prominent in most cases, flared with margin slightly ex- NUMBER 18 103 1 50/jm 1 278 279 282 283 FIGURES 274-283.—Kinorhynchus stenopygus, new species: 274, neck and trunk segments, ventral view, holotypic male (USNM 70026); 275, same, dorsal view; 276, same, segments 2-4, lateral half, ventral view; 277, same, dorsal view; 278, same, segments 12, 13, lateral half, ventral view; 279, same, dorsal view; 280, segments 2-4, lateral half, ventral view, allotypic female (USNM 70027); 281, same, dorsal view; 282, same, segments 12, 13, lateral half, ventral view; 283, same, dorsal view. 289 FIGURES 284-289.—Kinorhynchus stenopygus, new species: 284, ventral view, segments 2-4; 285, ventral view, segments 7, 8; 286, ventral view, segments 11-13; 287, dorsal view, segments 2-4; 288, dorsal view, segments 7, 8; 289, dorsal view, segments 11-13. (Interference contrast photographs all with same scale (in /xm) as shown in Figure 284; MP = middorsal process.) FIGURES 290-295.—Kinorhynchus stenopygus, new species: 290, episternal plate, segment 3, male; 291, adhesive tube, segment 4, male; 292, enlargement of Figure 291; 293, segments 7, 8, lateral half, ventral view, male; 294, segments 11, 12, lateral half, ventral view, male; 295, segments 12, 13, lateral half, ventral view, female. (SEM photographs, each to scale (in /xm) indicated; CS = cuticular scale, CH = cuticular hair, SE = seta.) 106 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES NUMBER 18 107 TABLE 20.—Measurements (/im) and indices (%) for Kinorhynchus stenopygus adults (see "Methods" for character abbreviations) Characte Standard Standard Coefficient r Number Range Mean deviation error of variability TL 6 4 520-603 556.3 38.5 19.2 6.9 2 9 509-592 543.9 32.8 10.9 6.0 62 13 509-603 545.7 34.2 9.9 6.3 SW 6 4 100-104 103.0 2.0 1.0 1.9 2 9 96-112 107.1 6.9 2.3 6.4 62 13 96-112 106.0 6.3 1.8 5.9 SW/TL 6 4 16.6-20.0 18.6 1.5 0.8 8.3 2 9 17.8-22.0 19.7 1.8 0.6 9.0 62 13 16.6-22.0 19.5 1.8 0.5 9.1 MSW-8 6 4 132-152 145.0 8.9 4.4 6.1 2 9 140-148 145.3 2.8 0.9 2.0 62 13 132-152 144.7 4.8 1.4 3.3 MSW/TL 6 4 24.5-27.9 26.1 1.5 0.7 5.7 2 9 24.7-28.8 26.6 1.6 0.5 5.9 62 13 24.5-28.8 26.4 1.6 0.5 5.9 tended posteriorly at midline (Figure 278). Three pairs of penile spines at anterolateral margins of sternal plates in males (Figure 278), simple gon- opores present in females. Males differ from females by the presence of 3 pairs of penile spines at anterolateral margins of terminal sternal plates, presence of 2 unmodified lateral setae on segment 12, presence of adhesive tubes on sternal plates of segment 4, even lateral margin on terminal tergal plate, and slightly narower trunk segments. Morphometric data for adults of Kinorhynchus stenopygus, new species, are shown in Table 20. HOLOTYPE.—Adult male TL 603 /xm (Figures 274-279, 284-289), Twin Cays, sta RH 444, Be- lize (16°50.0'N, 88°06.0'W), 8 Apr 1977, col. R.P. Higgins, USNM 70026. ALLOTYPE.—Adult female, TL 572 /xm (Figures FIGURES 296-301.—Kinorhynchus stenopygus, new species: 296, cross-sectional view of broken trunk, male; 297, posterior margins of segments 4, 5, enlarged from Figure 296; 298, segments 3, 4, dorsal view; 299, middorsal marginal process, segment 3; 300, segment 9, dorsal view; 301, middorsal marginal process, segment 10. (SEM photographs, each to scale (in /tm) indicated; PP = process pore, SS = sensory spot.) 280-283), other data as for holotype, USNM 70027. PARATYPES.—Two adult males and 8 females, TL 509-582 /tm, other data as for holotype, USNM 70028; 1 male, TL 572 /xm, Twin Cays, sta RH 442, other data as for holotype, USNM 70029. REMARKS.—Like Kinorhynchus deirophorus and K. distentus, K. stenopygus has unusually small penul- timate and terminal segments. It differs from the former two species by its prominent middorsal marginal projections which are distinctly spinose, not unlike those of K. anomalus. In fact, the ter- minal segment of this latter species, with its slightly flaring margin, is also similar to this same segment of K. stenopygus. The cuticular ridges of the sternal plates of K stenopygus are similar to those of several other new species described from this same locality. Espe- cially notable are the semicircular ridges which resemble those of K. belizensis and K. erismatus. The lack of thin areas of cuticle at the anterior margins of the episternal and midsternal plates is thus far distinctive among the Belizian material, and the three small points positioned middorsally and laterally on the anterior margin of the tergal 108 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES plate of the first trunk segment are unique to the genus Kinorhynchus; similar tergal margins have been described for Pycnophyes ponticus, P. keilensis, and P. neapolitanus. ETYMOLOGY.—This species name is from the Greek steno (narrow) plus pygos (rump), which reflects the morphology of the terminal segments. Kinorhynchus trisetosus, new species FIGURES 302-335 DIAGNOSIS.—Trunk slightly tapered anteriorly and posteriorly; anterolateral tergal projections pointed, similar projections situated laterotermin- ally at the tergal-sternal junction of the penulti- mate segment, extending beyond terminal seg- ment; anterodorsal tergal margin denticulate; middorsal margin spinose processes on segments 3-11; 3 lateral setae on segment 12. DESCRIPTION.—Adults (Figures 302-335) trunk length 676-832 /xm; trunk slightly tapered ante- riorly and pcsteriorly; MSW-8 204-236/xm, 26.8- 32.5 percer of trunk length; SW 148-176 /xm, 20.0-25.5 percent of trunk length. Second seg- ment with 4 dorsal and 2 ventral even-margined placids (Figures 304, 319, 322). Trunk segments (Figures 302, 303) with spinose middorsal marginal projections on segments 3- 11. Setae, 20-22 /xm long, situated middorsally on segment 10, perispinally on tergal plates of segments 9-11, subdorsally on segments 3-10, dorsolaterally on segments 3-11 with single seta lateral to each dorsolateral seta of segment 3; 1 seta between dorsolateral and subdorsal setae on segments 5-7, 9, 10, and 2 setae between the dorsolateral and subdorsal setae of segment 8; lateral setae on segments 4-11 with 3 lateral setae on segment 12 of males, sometimes not evident in females (Figure 332); single lateral setae on ster- nal plates of segments 5, 6, 8-11, 2 such setae on segment 7; mesial setae on segments 4, 6-11. Series of parallel cuticular ridges and cuticular ridge broadly looping posteriorly on sternal plates of segment 3; more narrow, smaller cuticular loops on sternal plates of segments 5-8 (Figures 302, 332, 333). Perispinal sensory spots on ante- rior 3-4 tergal plates (Figure 324); subdorsal sensory spots on segments 3-5 with 2 additional sensory spots on tergal plates of segment 3 (Fig- ures 319, 321); dorsolateral sensory spots on seg- ments 6-11; several smaller sensory spots (which are difficult to see without SEM) on segment 10 and more randomly on other tergal plates; 2 sensory spots, 1 anterior (Figures 330, 331) and 1 posterior (Figure 332), on episternal plates of segment 3; single sensory spot on each sternal plate of segments 4-10, 12 (Figures 330, 331) with 2 ventral sensory spots on sternal plates of seg- ment 11 (Figure 334). Large oval muscle scars dorsolateral on first trunk segment (Figures 303, 305-309), more prominent on episternal plates; several triangular cuticular scars anteromesial to larger muscle scars, small oval cuticular scars subdorsal on segments 4-12 (Figure 303), small, round to oval scars on episternal plate of segment 3 and sternal plates of segments 3-12. Pachycycli well developed; no midventral thickenings ob- served. Cuticle with some elongate scales on tergal plates and lateral margins of sternal plates, form- ing hairlike processes at tergal-sternal junctions (Figure 323); cuticular surface over large oval muscle scars of tergal plate 3 with striations (Fig- ure 322), short cuticular hairs appear to replace scaly cuticular surface on mesial portions of ster- nal plates, progressively more prominent on pos- terior segments. First trunk segment (segment 3) with pointed anterolateral tergal projections (Figures 304, 309, 319, 322); anterodorsal margin denticulate (Fig- ures 305, 309, 321). Muscle scars, setae, and sen- sory spots as previously noted; no areas of thin cuticle visible by light microscopy at anterior margins of episternal or midsternal plates al- though these areas are set apart by fine superficial sculpturing when viewed with SEM (Figure 330). Midsternal plate elongate-trapezoidal, lateral margins straight, 131 /xm long, 86 /xm basal width, 30 /xm width at apex; apex slightly expanded at margin only. Sternal plates of segment 4 of males with ad- hesive tubes 48 /xm long (Figure 304); females lack adhesive tubes but have 2 setae, 1 on either NUMBER 18 109 T 50 um FIGURES 302-311.—Kinorhynchus trisetosus, new species: 302, neck and trunk segments, ventral view, holotypic male (USNM, 70030); 303, same, dorsal view; 304, same, segments 2-4, lateral half, ventral view; 305, same, dorsal view; 306, same, segments 12, 13, lateral half, ventral view; 307, same, dorsal view; 308, segments 2-4, lateral half, ventral view, allotypic female (USNM 70031); 309, same, dorsal view; 310, same, segments 12, 13, lateral half, ventral view; 311, same, dorsal view. 110 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 317 FIGURES 312-317.—Kinorhynchus trisetosus, new species: 312, segments 3, 4, ventral view; 313, segments 7, 8, ventral view; 314, segments 11-13, ventral view; 315, segments 3, 4, dorsal view; 316, segments 7, 8, dorsal view; 317, segments 11-13, dorsal view. (Interference contrast photographs all with same scale (in /im) as shown in Figure 312.) FIGURES 318-323.—Kinorhynchus trisetosus, new species: 318, trunk segments, lateral view; 319, neck and first trunk segment, dorsal view; 320, neck with trichoscalids, dorsal view; 321, anterolateral surface of first trunk segment; 322, neck and first trunk segment showing closed position, anterior view; 323, tergal-sternal junction. (SEM photographs, each to scale (in /xm) indicated; TR = trichoscalid.) FIGURES 324-329.—Kinorhynchus trisetosus, new species: 324, segment 4, middorsal marginal protuberance; 325, same, segment 9; 326, same, segment 10; 327, portion of intersegmental area from Figure 326 (enlarged); 328, same, segment 10 (enlarged); 329, tip of hollow seta. (SEM photographs, each to scale (in /im) indicated.) FIGURES 330-335.—Kinorhynchus trisetosus, new species: 330, segment 3, anterolateral view, episternal plate (female); 331, sensory spot, episternal plate (enlarged from Figure 330); 332, segment 4, sternal plate (female); 333, segments 7, 8, sternal plates, right half (female); 334, segment 12, sternal plates (female); 335, terminal margin with exposed penile spine. (SEM photographs, each to scale (in /im) indicated.) 114 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES TABLE 21.—Measurements (/im) and indices (%) for Kinorhynchus trisetosus adults (see "Methods" for character abbreviations) Standard Standard Coefficient Character Number Range Mean deviation error of variability TL 6 22 697-801 757.8 33.3 7.1 4.4 2 13 676-832 760.9 39.1 10.8 5.1 62 35 676-832 758.9 35.0 5.9 4.6 SW 6 22 148-168 161.3 4.8 1.0 3.0 2 13 156-176 167.1 6.8 1.9 4.1 62 35 148-176 163.4 6.2 1.1 3.8 SW/TL 6 22 20.0-23.0 21.3 0.9 0.2 4.3 2 13 20.0-25.5 22.0 1.5 0.4 6.9 62 35 20.0-25.5 21.6 1.2 0.2 5.6 MSW-8 6 22 204-236 216.7 9.4 2.0 4.3 2 13 212-236 221.4 5.9 1.6 2.6 62 35 204-236 218.5 8.5 1.4 3.9 MSW/TL 6 22 26.8-30.3 28.6 1.3 0.3 4.6 2 13 27.7-32.5 29.1 1.3 0.4 4.3 62 35 26.8-32.5 28.8 1.3 0.2 4.6 side of each adhesive tube site (Figures 308, 330). Several cuticular ridges with posterior loops pres- ent on sternal plates. Segments 5-11 similar with exceptions as previously noted. Segment 12 ta- pered with slightly uneven posterior tergal and sternal margins (Figures 306, 311, 334); 3 lateral setae present in males, more variable in females but usually indicated by setal pores in cuticle. Lateroterminal margins projecting beyond mar- gin of terminal segment. Terminal tergal segment with minute bulbous protuberance at lateroterminal margin, otherwise with even margin. Sternal plates even, generally with margins paralleling that of terminal tergal plate (Figures 306, 310). Three pairs of penile spines at anterolateral margins of terminal sternal plates in males (Figures 306, 335); single gonopore in females (Figure 310). Males differ from females by presence of 3 pairs of penile spines at anterolateral margins of ter- minal sternal plates, presence of adhesive tube on each sternal plate of segment 4, slightly narrower trunk, and some variation in sensory spots and setae. Morphometric data for adults of Kinorhynchus trisetosus are shown in Table 21. HOLOTYPE.—Adult male, TL 790 /xm (Figures 302-307, 312-317), Twin Cays, sta RH 442, Be- lize (16°50.0'N, 88°06.0'W), 8 Apr 1977, col. R.P. Higgins, USNM 70030. ALLOTYPE.—Adult female, TL 754 /xm (Figures 308-311), other data as for holotype, USNM 70031. PARATYPES.—Seventeen males and 9 females, TL 676-832 /xm, other data as for holotype, USNM 70032; 4 males and 3 females, TL 749- 801 /xm, Twin Cays, sta RH 443, other data as for holotype, USNM 70033. DISCUSSION.—Kinorhynchus trisetosus is the largest and most abundant of the kinorhynchs in the area of Carrie Bow Cay. On the basis of its size alone, it can easily be distinguished from other species. Because of the character of the extended- lateral margin of the penultimate segment which often exceeds the terminal margin of the last segment, K trisetosus most closely resembles K. mainensis. Several other characters are similar, including the bulbous projection situated postero- laterally on the terminal tergal plate, the general shape and dimensions of the midsternal plate, the setae of the episternal plate, and the relatively large number of sensory spots. NUMBER 18 115 The episternal and midsternal plates of K. mainensis have distinct areas of thin, sculptured cuticle readily apparent in all specimens when viewed by light microscopy; this is not the case in K. trisetosus. Kinorhynchus mainensis does not have a denticulate margin on the tergal plate of the first trunk segment nor does it have middorsal spinous marginal projections on the margins of most trunk segments as in K. trisetosus. Furthermore, those spinose projections are much smaller, less distinctive in K. mainensis. This latter species has lateral setae on segments 3, 4, 6, 8, 10 and two on 12 as opposed to 4-11 and three on 12 (occasion- ally absent or less apparent in females) in the new species. Altogether, the two species are distinct. Kinorhynchus giganteus, heretofore the largest of the genus Kinorhynchus, is easily distinguished from K. trisetosus on the basis of the crescent-shaped cuticular scars on the sternal plates of K. giganteus and its bi- or trifid lateroterminal projections and lateral setation pattern differences. ETYMOLOGY.—The name of this species is from the Latin tres (three) plus seta (bristle), reflecting the three lateral setae condition of the penulti- mate segment. Discussion of Kinorhynchus As a kind of progress report on his preparation of Monographic der Echinodera, Carl Zelinka, in 1895, demonstrated his illustrations to an audi- ence of the Deutsches Zoologisches Geselschaft. The following year, when the proceedings were published (Zelinka, 1896), several new taxa and their descriptions were in print for the first time. Included in this publication was the family Trachydemidae but not its nominal genus Trachydemus. In 1907, under the title Zur Kenntnis der Echinoderen, Zelinka (1907:135) established the genus Trachydemus and the "larval genus" Lepto- demus that in part, along with the "larval genus" Centrophyes, contained juveniles that later com- prised the "larval genus" Hyalophyes. Specimens of the latter genus are juveniles of Pycnophyes. Other forms of Leptodemus are juveniles of Trachydemus. Thus, at that time, the order Homalorhagida consisted of two families: Pyc- nophyidae with its nominal genus Pycnophyes and three "larval genera" (synonyms) Hyalophyes, Centrophyes, and Leptodemus (part), and Trachy- demidae with its nominal genus Trachydemus and "larval genus" (synonym) Leptodemus (part). Remane (1936:352, 353) first suggested that the two homalorhagid families be united; Lang (1949:11-14) formalized this union. It was not until 1974 that Sheremetevskij discovered that the name "Trachydemus'''' and its junior synonym Leptodemus were preoccupied by Trachydemus Chevrolat, 1873 (Coleoptera), and Leptodemus Reuter, 1900 (Hemiptera), respectively. Conse- quently, Sheremetevskij substituted the generic name Kinorhynchus Sheremetevskij, 1974, for Trachydemus Zelinka, 1907. For many years, the only described species of adult Kinorhynchus was K. giganteus. This species was found on both the northern and Mediterra- nean coasts of Europe (Table 22). Nine other species, probably of Kinorhynchus, include: Lepto- demus acercus and L. metschnikowii (considered Pyc- nophyes acercus by Bacescu, 1968), L. dubius, L. forficulus, L. parvulus, L. perlatus, and L. vitreus (considered Pycnophyes by Remane, 1929), L. for- ceps, and L. serratus. As noted previously in my discussion of Pycnophyes, I prefer to assume that without greater evidence than currently avail- able, these species should be in the genus Kino- rhynchus, although it is doubtful that any will be united with a given adult. The second species of Kinorhynchus based on an adult was K. mainensis, which was redescribed from material collected at the type-locality, Mt. Desert Island, Maine, USA (Higgins, 1965). The third such species of Kinorhynchus was K. spinosus from the Falkland Islands (Lang, 1949). The next species, K anomalus, was collected from the coast of Chile (Lang, 1953). Kinorhynchus ilyocryptus and K cataphractus were described from the San Juan Archipelago, Washington, USA (Higgins, 1961). Kinorhynchus langi was described from Beaufort, North Carolina, USA (Higgins, 1964b). The most recently published species, K. paraneapolitanus, was described by Sheremetevskij (1974) from the Black Sea. 116 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES Considering the number of collections made over the past hundred or so years, it is noteworthy that only eight species of Kinorhynchus (based on adults) have been described compared to the closely related genus Pycnophyes with its 18 species published on the basis of adults. More important, perhaps, is the frequency by which Pycnophyes (Table 15) is encountered as compared to that of Kinorhynchus (see Table 22). The genus Kinorhyn- chus, like Pycnophyes, is found in a wide variety of habitats but appears to have a limited distribu- tion in the world oceans (Figure 336), and very little can be construed from the published distri- bution records. Many authors have commented on the validity of maintaining the separation of the two genera of Pycnophyidae. Remane (1936) was the first to question the propriety of the two genera, while Lang (1949) defended the continual separation of the two genera, and this has persisted to date. The only character separating these two genera is the presence of lateral terminal spines in Pyc- nophyes. So far, whenever an appendage appears in the equivalent position in the genus Kinorhyn- chus, it appears to be an unarticulated extension of the lateral or lateroterminal margin of the terminal tergal plate. This protuberance has been described as a bifid (male) or flaring (female) portion of the tergal plate in K. giganteus (Zelinka, 1928), as a bulbous projection in K. mainensis (Blake, 1930), or as stunted lateral terminal spines in K. spinosus. Until it can be demonstrated that these structures are articulated and attached to muscles as in the case of all known species of Pycnophyes, the genus Kinorhynchus should, in my opinion, remain distinct. To date, there has been no concerted effort to conduct a contemporary comparative study of the two genera. In addition to developing culture methods to enhance the study of the life histories of members of the two genera, ultrastructural studies of various organs, such as adhesive glands, sensory spots, setae, and careful observations on the detailed structure of the head should be pur- sued for additional data to better understand their relationships. Meanwhile, another important task remains: that of carefully redescribing most of the previ- ously described species, including K. giganteus, in order to more carefully assess the number and position of characters such as sensory spots and setae. In several species such as K. spinosus, only lateral setae were noticed, and no sensory spots were evident. Certainly more detailed illustration of the trunk segments of this and other species, including K. ilyocryptus, K. cataphractus, and K. langi originally described by me, will be necessary in the near future. In my opinion, however, only Kinorhynchus paraneapolitanus is so poorly described and illustrated as to prevent its accurate identi- fication. Species Distribution and Richness Prior to this publication, 119 papers by 52 different authors have included distribution rec- ords. Of these, 80 papers have included species identifications. These 80 papers have involved, as best as I can determine, 182 samples, stations, or localities. Regardless of the limits of the collecting unit involved, the resulting data give one the overwhelming impression that rarely does one encounter more than one or two genera or more than three or four species within the collecting units (Table 23). The greatest recorded species richness was re- ported from Naples, Italy (Figure 337), by Ze- linka (1928). From his Pallazzo Domano station (sta 5) at 35 m depth, Zelinka found seven species representing three genera (Figure 338). Alto- gether, Zelinka sampled 18 stations, 13 of which produced at least one kinorhynch. His habitats ranged from algae to mud, or combinations of the two, and depths of 1-65 meters. Data con- cerning the specimens from seven of the produc- tive stations are incorporated into Figure 338 to illustrate both species distribution and richness within the study area (about 14 km of coastal water) in the Bay of Naples. Relative abundance indications in the figure are not based on real numbers, but on Zelinka's commentary. Zelinka's study of 19 stations in the Gulf of NUMBER 18 117 TABLE 22.—Distribution of Kinorhynchus (see Figure 336); type-localities are in italics) Species FAO sea area Locality Authority anomalous PSW W. Estero Reconcavi, Chile Lang, 1953 PSW W. Canal Moraleda, Chile Lang, 1953 apotomus, ASW Twin Cays, Belize Higgins, this study new species belizensis, ASW Twin Cays, Belize Higgins, this study new species cataphractus INE Sucia Is. (Wash.), USA Higgins, 1961 INE Orcas Is. (Wash.), USA Higgins, 1961 INE Stuart Is. (Wash.), USA Higgins, 1961 deirophorus, ASW Twin Cays, Belize Higgins, this study new species distentus, ASW Twin Cays, Belize Higgins, this study new species erismatus, ASW Twin Cays, Belize Higgins, this study new species giganteus ANE Kiel, FRG Zelinka, 1928 MED Naples, Italy Zelinka, 1928 MED Trieste, Italy Zelinka, 1928 ilyocryptus INE Orcas Is. (Wash.), USA Higgins, 1961 INE Sucia Is. (Wash.), USA Higgins, 1961 INE Stuart Is. (Wash.), USA Higgins, 1961 INE Tomales Bay (Calif), USA Higgins, 1964b langi ANW Beaufort (N.C.), USA Higgins, 1964b ANW Marineland (Fla.), USA Higgins, 1964b mainensis ANW ML Desert IS. (Me.), USA Blake, 1930; Higgins, 1965 ANW Buzzards Bay (Mass.), USA Wieser, 1960; Higgins, 1965 ANW Lewes (Del.), USA Higgins, 1965 paraneapolitanus MED Lazarevskoye, USSR Sheremetevskij, 1974 spinosus PSW Barkley Sound, Falkland Is. Lang, 1949 stenopygus, ASW Twin Cays, Belize Higgins, this study new species trisetosus, ASW Twin Cays, Belize Higgins, this study new species species INW Kasado Is., Japan Sudzuki 1976a,b species MED Bulgaria Marino\ r, 1964 Trieste (Figures 339, 340) produced similar re- sults. Here again, a maximum of seven species (three genera) were found at stations 2, 3, and 9; 10 species representing four genera were found in this overall area, which included the smaller Bay of Piran south of Trieste. The most meaningful data came from samples taken along a seven-mile transect from station 1 (17 m) to station 4 (24 m) off the coast of Barcola (Figure 339). Two of Zelinka's Trieste stations, stations 2 and 9, had the same seven species present. One additional station, station 7, had seven species; six of which were the same as those found in stations 2 and 9. Zelinka's studies in these two Mediterranean lo- calities, the Bay of Naples and Gulf of Triste, account for most of the highest records of species richness shown in Table 23. Unlike the majority of these records, Zelinka's work was the product of research directed toward only the kinorhynch fauna of his study sites, which may account for some of his results; however, where the few other similar intensive studies of a given locality have 118 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES FIGURE 336.—Distribution of Kinorhynchus (solid circles indicate records of named species, stippled circles indicate genus records only). TABLE 23.—Summary of frequency of species and genera reports per sample site (* = this study) Maximum Maximum number of Number of number of Number of genera/site sample sites species/site sample sites 1 127 1 119 2 34 2 25 3 17 3 17 4 4+1* 4 5 6 7 8 13 10 5 3 2 1 1* been conducted (Higgins, 1960, 1961, 1964b; Ny- holm, 1947b; Wieser, 1960; Mclntyre, 1962, 1964; Sheremetevskij, 1974; Bacescu, 1968), the results have indicated that the meiobenthic com- munity generally accommodates three genera or less with four species or less, if the samples are restricted to subtidal sediments only. A series of collections made in the San Juan Archipelago (Figure 341), Washington, USA (Higgins, I960, 1961; Kozloff, 1972), illustrates a typical com- munity structure of five species representing three genera (Figure 342). Considering the previous records of relatively low kinorhynch species richness within given re- stricted localities, most of them at temperate lat- itudes, the results of the limited sampling in the Carrie Bow Cay study area are exceptional al- though, perhaps, in keeping with our general concept of high species richness in tropical eco- systems. Although this study was not designed to support such ecological conclusions, possibly the greater environmental heterogeneity in this trop- ical reef ecosystem is the reason for the accom- modation of the unprecedented number of kino- rhynch species found. Eighteen new species representing four genera, one of these new, were the product of a single dredge sample at each of six stations. The two richest stations included RH 442, located in the mangrove channel system of Twin Cays, and RH 444, located 1 km to the southwest in a typical area of seagrass. Each station produced 13 species (Figure 343). The mangrove station with its very fine gray-brown mud at a depth of 1-2 meters had 13 species representing three genera: Kinor- NUMBER 18 12 @ ® 13 )4 ®3 O ®5 ®6 8 7b N7a % >® o 16°E Gulf of Naples Yugoslavia o 1 i i km O 40 N Tyrrhenian Sea "\ VJ FIGURE 337.—Map showing Gulf of Naples sample stations (Zelinka, 1928) (open circles indicate negative sample, circles with solid inner circles indicate selected stations referred to in Figure 338, circles with open inner circles indicate other stations where kinorhynchs were collected; area of larger map located on inset by arrow). 120 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES Relative Abundance Taxa 3 ■c Oco oS. D> H Abundant O Common c j Uncommon V) V) (O s« Q> lat ■fccB § •OS o Q)C o = op CO 0"D S2S C.to ■C'3" cpo- CB OTD <0 UJ Taxa SEDIMENT Station 1 19m Station 2 19 m Station 3 22 m Station 4 24 m Station 9 17m ALGAE Station 13 3m ft s (S) 1 mm FIGURE 340.—Diagrammatic representation of selected Gulf of Trieste stations (Zelinka, 1928), their species composition and relative abundance. All stations were sampled within the same month, and the results are assumed to be the product of a single sample. NUMBER 18 123 Reid Harbor c v*"^?^ /? V Pacific Ocean Y%X Canada \> ST. vl V *Zf USA FIGURE 341.—Map showing islands of the San Juan Archipelago (Washington, USA) sample stations (Higgins, 1960, 1961; Kozloff, 1972) (area of larger map located on inset by arrow). 124 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES o O Rare Abundant Common o Uncommon Relative Abundance Taxa Sediment Station 1 32 m 1 mm Station 2 30 m Station 3 10 m Station 4 Intertidal FIGURE 342.—Diagrammatic representation of San Juan Ar- chipelago stations (Higgins, 1960, 1961; Kozloff, 1972), their species composition and relative abundance. Stations 1-3 were sampled during the same month, station 4 was sampled at various periods of time, not in the same year. hynchus, Pycnophyes, and Echinoderes. The seagrass station, RH 444, had 13 species representing four genera: Kinorhynchus, Pycnophyes, Paracentrophyes, and Echinoderes. Eight of the 18 species were found at both the mangrove and seagrass stations noted above (Fig- ure 343), seven of these were also found at the intermediate station, RH 443, a less stable site as indicated by the slightly coarser sediment and less dense, more patchy Thalassia growth. Within the group of species found at both the mangrove (RH 442) and seagrass (RH 444) sites, their relative abundance indicates that while they may be found in both habitats, one habitat apparently is preferred over the other in at least a few cases. This is illustrated by Kinorhynchus erismatus and K. stenopygus, which are more abundant in the sea- grass habitat, whereas K trisetosus and K. belizensis and possibly Pycnophyes longcornis appear to prefer the mangrove habitat. Only two of the five species of Echinoderes, E. horni and E. wallaceae, were found in any of the three stations along the reef transect where conditions reflected the higher energy, less stable sediment of this ecosystem. Nine (representing all four genera) of the area's 18 species were found at only one station. These included Kinorhynchus deirophorus, K. distentus, K. apotomus, Pycnophyes iniorhaptus, P. corrugatus, P. ec- phantor, Paracentrophyes praedictus, Echinoderes trun- catus, and E. abbreviatus. Although the evidence is limited, it suggests that these species are probably restricted to one of the two habitats, mangrove or seagrass. No species was restricted to the inter- mediate, less stable habitat (sta RH 443) between the two other stations. Although Echinoderes wallaceae was not present in the sand rubble sample, and E. horni was absent from the reef trough sample, I suspect both E. wallaceae and E. horni would have been found in these stations if a larger number of samples could have been taken. Of the five local species in the genus, E. wallaceae and E. horni have the longest lateral terminal spines in relation to the trunk length. Both E. truncatus and E. abbreviatus were found at a single station each. The latter species, with its relatively short lateral terminal spines as con- trasted with E. wallaceae and E. horni with their very long lateral terminal spines, would appear to support the hypothesis (Higgins, 1967) that short-spined species are adapted to fine sediments and longer spines to coarser sediments. snieiAejqqe GJEHH&/ j o co i 6 lujoq GSBEBSCI j smejofjadw! IfliillLLfgQ tmrminQ; snjeounjj sajapouiqo3 HfttHtfrQ E smoipaejd E saAudojjuaoejed o o jotuBL/doa SIUJOOIBUO) O snteBnjJoo snieuiBjeuua snideqjoiui saAudouoAd O *-«»«» afjte* am snuiotode snBAdouais sntuaisfp snjoudojiap snieuisua C^Fj. snuouAuJOUM K-T snsoiasjJi t 3 tm t, OJ V m r, OJ o u c nj ft ■J-J C OJ a N JD tf u CQ > OJ *^ X. (Ti <— 0 OJ ion -a G r3 rt c 0 C o; --■ u n n. a F u 0 o u - = ~ - t- br> Kl Q I i on -v zn a ei z. _- CO o c CD CO *= C .2 R QC*3- LLI > C\J co "* o "* Z QC o c c axa /IANG Statio 1-2m .AGO Statio 3.3m Statio 3m 2 E CO C\J = E = E 126 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES Phylogenetic Relationships Apart from the phylogenetic relationships dis- cussed in connection with Paracentrophyes praedic- tus, several observations should be made concern- ing other kincrhych taxa found in Belize. Perhaps the most unexpected find was the seven new species of Kinorhynchus. Only once have more than two species of this genus been found in a single station or even within a several hundred square kilometer area (Higgins, 1960, 1961). The previ- ous total number of species of this genus was only eight; seven new species from one area has nearly doubled the number of known species of this genus. Excluding any relationship with Kinorhynchus paraneapolitanus, which is so poorly described that no comparison is practical, there seems to be no close phylogenetic relationships between the Be- lizian material and the remaining species; how- ever, two pairs of Belizian species show distinct relationships that should be noted. The first two species that are remarkably sim- ilar are Kinorhynchus deirophorus and K distentus. Both have reduced penultimate segments, a char- acter also shared with K. stenopygus but with no other species in the genus. The structure of the first trunk segment is a second important shared character. Both species have a similar pattern of thin cuticular areas, two on each episternal plate and two on each midsternal plate, and the shape of the midsternal plates is nearly identical. Some similarity can be found between this condition and that of A', giganteus from both the north and south coasts of Europe, but this latter species appears to have three thin cuticular areas on the midsternal plate rather than two. Therefore, this character state in the species from Belize appears to be unique. Likewise the tergal plates of the first trunk segment of A', distentus and K. deirophorus are similar, especially in the pattern of cuticular ridges and the close proximity of the two promi- nent sensory spots on either side of the dorsal midline. Other, less obvious, shared character states include the structure of the pachycycli of the sternal plates of segments 10 and 11. This is especially noticeable when one compares the mid- ventral thickenings of the two species. Also, the cuticular ridge pattern of the sternal plates of the fourth segment appear remarkably similar. Nei- ther species has middorsal marginal processes. The second pair of species that share character states, some of which are unique to the genus, include K. erismatus and K. belizensis. The most obvious, as well as uniquely, shared character state is the structural element between the lateral margins and pachycycli of the sternal plates of segments 4-10 or 11. This "buttressing" feature has not been seen in other species of kinorhynch. A second shared unique character state is the lack of any lateral setae on the penultimate segment. In fact, the margin appears to have two cuticular interruptions in the area where setae are usually found. Some additional similarities include the pattern of muscle scars on the tergal plates of the first trunk segment and some similarities between the episternal plate structure and cuticular ridges of the sternal plates. Kinorhynchus deirophorus and A. distentus appear to occupy separate habitats (Figure 343); how- ever, both A. erismatus and A. beliziensis are found in all three Twin Cays sites, although the latter species appears to be distinctly more abundant in the mangrove habitat (Figure 393). These two pairs of Kinorhynchus species appear to have evolved, each from the same common ancestor, which may or may not include an extant species, or one from the other, but only one pair appears to have persisted in both seagrass and mangrove habitats. Within the genus Pycnophyes, two of the five Belizian species lack the adhesive tubes found in all other males of the genus except for P. egyptensis. The two local species are P. longicornis and P. ecphantor. Aside from the lack of adhesive tubes, the only other notable similarities between the two species include the arrangement of setae and sensory spots on the sternal plates, the denticulate margin of the tergal plate of segment 3 (common to other species as well), and the length and shape of the lateral terminal spines. Pycnophyes corrugatus has a unique series of ridges near the lateral margins of the sternal plates. This NUMBER 18 127 character state is slightly suggestive of a similar pattern of striations in this same area in P. denta- tus. Pycnophyes corrugatus, with regard to this char- acter state, is most similar to Kinorhynchus apotomus. Moreover, the two species share several other characters such as a general similarity of seta distribution, a single, thin cuticular area on each episternal and midsternal plate, a denticulate anterodorsal margin of the first trunk segment, and some similar cuticular ridges on the same tergal region. Both are restricted to the more stable Thalassia station (RH 444) of the Twin Cays area. In addition to the previously mentioned unique character states found in these Belizian species, the unusual patches of pores near the lateral margins of the sternal plate of A. iniorhaptus should also be mentioned. This character is found in no other known kinorhynch. The species of Echinoderes from Belize show no clear relationships to each other as far as I can determine. Perhaps the most noteworthy relation- ship between any known species of Echinoderes from Belize to any other member of this genus is that between E. abbreviatus and E. brevicaudatus from the Egyptian Red Sea. Both have notably short lateral terminal spines, and both have been found in coral-derived fine sediments. Literature Cited Abe, Y. 1930. Das Vorkommen von Echinoderes in den japan- ischen Gewassern. Journal of Science of the Hiroshima University, series B, l(l):39-44, 2 figures, 1 plate. Ankar, S., and R. Elmgren 1976. 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