Redescription of

Echinoderes dujardinii (Kinorhyncha)

with Descriptions of

Closely Related Species

ROBERT R. HIGGINS

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 248
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SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 248

Redescription of

Echinoderes dujardinii (Kinorhyncha)

with Descriptions of

Closely Related Species

Robert P. Higgins

SMITHSONIAN INSTITUTION PRESS

City of Washington

1977
ABSTRAC T

Higgins, Robert P. Redescription of Echinoderes dujardinii (Kinorhyncha)

with Descriptions of Closely Related Species. Smithsonian Contributions to

Zoology, number 248, 26 pages, 31 figures, 2 tables, 1977.—A key to the adults

of the genus Echinoderes is followed by a redescription of E. dujardinii from

both adult and juvenile specimens. North American records for E. dujardinii

are corrected by the description of E. kozloffi, new species. Echinoderes pennaki,

a species sympatric with E. kozloffi, is redescribed, and the latter's sibling species,

E. pacificus, is discussed. Synonomies and distribution records are given for

species mentioned.

OFFICIAL PUBUCATION DATE is handstamped in a limited number of initial copies and is recorded

in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: The coral Montastrea

cavernosa (Linnaeus).

Library of Congress Cataloging in Publication Data

Higgins, Robert P.

Redescription of Echinoderes dujardinii (Kinorhyncha) with descriptions of closely related

species.

(Smithsonian contributions to zoology ; no. 248)

Bibliography: p.

Supt. of Docs, no.: SI 1.27:248

I. Echinoderes dujardinii. 2. Echinoderes. 3. Kinorhyncha—Classification. I. Title: Redescription of Echinoderes dujardinii (Kinorhyncha) ... . II. Series: Smithsonian Institution.

Smithsonian contributions to zoology ; no. 248.

QL1.S54 no. 248 [QL391.A2] 591'.08s [595M83] 76-608300
Contents

Page

Introduction 1

Methods 2

Acknowledgments 3

Key to Adults of the Genus Echinoderes 3

Echinoderes dujardinii Claparede 4

Echinoderes kozloffi, new species 13

Echinoderes pennaki Higgins 18

Echinoderes pacificus Schmidt 21

Echinoderes brevicaudattis, substitute name for E. brevispinosus Higgins .. 21

Literature Cited 24

Redescription of

Echinoderes dujardinii (Kinorhyncha)

with Descriptions of

Closely Related Species

Robert P. Higgins

Introduction

The first published observation of a kinorhynch

is that of the French naturalist F£lix Dujardin

(1851), who based his account on specimens found

while examining material washed from algae collected near St. Malo on the coast of Normandy in

1841. In a manner not uncommon throughout the

history of the study of meiofauna, Dujardin was

unable to place the new invertebrate in an existing

higher taxonomic category although he recognized

its morphological affinities with acanthocephalans,

rotifers, sipunculids, nematodes, tardigrades, and

various crustaceans ("copepods without legs"). Ten

years after his discovery, Dujardin (1851) introduced the first kinorhynch as "a tiny marine animal, I'Echinodere, constituting an intermediate

form between the crustaceans and worms." A few

years later Leuckart (1854) noted that he had seen

"I'Echinodere" at Helgoland in 1846 but had assumed that it was a dipteran larva.

"L'Echinodere" became Echinoderes dujardinii

Claparede, 1863 when the latter author described

specimens from St. Vaast la Hougue, not far from

St. Malo where Dujardin made his discovery. The

following year, Gosse (1864), apparently unaware

Robert P. Higgins, Smithsonian Oceanographic Sorting Center, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560.

of Claparede's publication, also described this species from the same locality, naming it Echinodera

dujardini.

These descriptions suffered from the authors' lack

of experience with the taxon, as have many subsequent ones, including my own. Despite Zelinka's

(1928) more comprehensive description of E. dujardinii in his Monographic der Echinodera, certain taxonomic ambiguities have persisted. The

lack of preserved specimens, especially types, has

compounded the problem; consequently, the reported distribution of E. dujardinii, as well as

other species, must be considered with due caution.

Since it was described, E. dujardinii has been reported from 26 localities including the northern

and southern coasts of Europe, the Black Sea,

Canary Islands, Japan, and the northwestern coast

of the United States.

No specimens are available to confirm Chitwood's (1964a) report of E. dujardinii from Tomales

Bay, California, one of the two northwestern U.S.

localities. Neither an unpublished photograph by

Chitwood (pers. comm.) of one of the specimens,

nor sketches of specimens collected at the same

locality and time (1960) by Dr. Tor G. Karling

(pers. comm.), indicate that the species from

Tomales Bay is E. dujardinii. Specimens sent to me

from San Juan Island, Washington, by Dr. Eugene

N. Kozloff, resembled E. dujardinii sufficiently to

1
2

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

justify the tentative use of this name (Kozloft,

1972; Merriman and Corwin, 1973); however, both

Kozloft and I expressed doubts that could be answered only by comparing the San Juan Island

specimens with specimens from either the typelocality or localities (Naples and Trieste) where

Zelinka (1928) obtained E. dujardinii from his

redescription. My collection from the latter two

localities failed to provide specimens of E. dujardinii but Kozloff, in 1973, was successful in collecting this species at Roscoff, not far from St. Vaast

la Hougue.

The purpose of this paper is to redescribe E. dujardinii, clarifying both the original description by

Glaparede (1863) and the redescription by Zelinka

(1928). In addition, this paper will correct the distribution record inasmuch as the San Juan Island

specimens, reported as E. dujardinii, constitute a

new species described below. Because of the cooccurrence of the new species with E. pennaki

Higgins, 1960, the description of which reflects

my own inexperience at the time, I shall redescribe

it as well. The description of E. pacificus Schmidt,

1974 is included because it closely resembles the

new species from San Juan Island. Finally, while

studying the taxonomic history of E. dujardinii, I

discovered that a name I proposed for a species

from the Red Sea was preoccupied; therefore, a

substitute name is offered for E. brevispinosus Higgins, 1966a.

METHODS.—Specimens were preserved in 70 percent ethyl alcohol, 5 percent formalin, or Duboscq

and Brasils fluid, and then transferred to a 70

percent ethyl alcohol-5 percent glycerin solution

that was allowed to evaporate to glycerin. Most

specimens were removed from the glycerin and individually placed in 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 specimen 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 exoskeleton.

A disadvantage of the 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 reducing, by about 30 percent, the amount of chloral hydrate used in the

medium and by sealing the preparation with Murrayite soon after the fluid has solidified. In this

series of preparations, some specimens were remounted in glycerin once they had been oriented

dorsoventrally. This is a procedure that should be

used only when an adequate series of specimens are

available, since in transferring them many dorsal

spines are often broken. A few specimens were

mounted in glycerin without first mounting them

in Hoyer's medium. In all instances, coverslips were

sealed with Murrayite.

Each specimen was studied with the use of Zeiss

differential interference contrast optics and meristically analyzed. The resulting data are expressed

in a standard format of abbreviations and terminology (Higgins, 1967, 1969a). Measurements are

given in microns (urn); ratios (i.e., SW/TL) are

expressed in percent of the total length (TL) measured on the midline, from the anterior margin of

segment 3 (first trunk segment) to the posterior

margin of segment 13, exclusive of spines. Maximum sternal width (MSW) is measured at the

anteroventral margin of the widest pair of sternal

plates as first encountered in measuring 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.

Middorsal spines (D), lateral spines (L), and

lateral accessory appendages (LA) are numbered by

segment and their cumulative mean length expressed by Dm, Lm, and LAm, respectively. Measurements are given for the lateral terminal spines

(LTS), lateral terminal accessory spines (LTAS),

midterminal spine (MTS), and penis spines (P) in

males. The first penis spine (P-l) is usually the

anteriormost of three such spines and is dorsally

displaced; the second (P-2) is usually the shortest

and often more truncate (probably the functional

penis); the third (P-3) is usually adjacent to the

second or slightly posterior to it. Both P-2 and P-3

are best observed in ventral aspect.

Several lateral spines appear to function as adhesive tubes. This study will introduce the adhesive

tubes of the fourth segment (L-4) as homologues

of other lateral spines. Appendages that function

as adhesive tubes will be noted in the appropriate

section of the text.
NUMBER 248

Specimens mentioned in this paper are deposited

in the National Museum of Natural History, Smithsonian Institution, under the catalog numbers of

the old United States National Museum (USNM).

ACKNOWLEDGMENTS.—I am grateful to Mr. John

C. Boykin, University of Washington, the first to

collect the new species from San Juan Island and

recognize its taxonomic problems; and to our mutual friend and colleague from this same institution, Dr. Eugene N. Kozloff, who accepted the

challenge of culturing the new species, and provided me with specimens both of the new species

and of E. dujardinii that have made this study

possible. I also acknowledge the cooperation of Dr.

Jean Merriman, California State College at Sonoma,

whose recent doctoral dissertation and publication

have contributed to the knowledge of the new

species from San Juan Island. The personal communications of Dr. Tor G. Karling, University of

Stockholm, and the late Dr. B. G. Chitwood were

helpful in this study. Dr. Peter Schmidt, Institute

fur Zoologie der RWTH, Aachen, generously provided type specimens of E. pacificus for my use.

Thanks also go to Prof. Peter Dohrn, Stazione

Zoologica, Naples and to Dr. Jose Stirn, Marine

Laboratory, Portoroz, Yugoslavia, for making their

facilities available to me. Drs. Raymond B. Manning

and Horton H. Hobbs, Jr., were generous in their

advice and consultation during the preparation of

this manuscript and, along with Dr. Richard S.

Houbrick, in the reading of the final draft. I also

wish to express my appreciation to Mrs. Eleanor

Goldsmith, Mrs. Martha Brewster, and Mrs. Marie

Wallace for their help in the preparation of this

manuscript. Finally, I am pleased to acknowledge

the Sumner Gerard Foundation and the Smithsonian Foreign Currency Program for their financial support.

Key to Adults of the Genus Echinoderes

1. Middorsal spines absent 2

Middorsal spines present 5

2. Lateral spines (except for terminal segment) absent

Echinoderes maxwelli (Omar-Cooper, 1957)

Lateral spines present 3

3. Lateral spines on segments 7, 10, and 12 only Echinoderes bengalensis (Timm, 1958)

Lateral spines absent on segment 12 4

4. Lateral spines on segments 7, 10, and 11 only Echinoderes caribiensis Kirsteuer, 1964

Lateral spines on segments 7 and 10 only, segments 3-4 expanded laterally

Echinoderes capitata (Zelinka, 1928)

5. Middorsal spines on segments 6-10 only 6

Middorsal spines with other arrangement IS

6. Lateral spines on segments 7-12 only 7

Lateral spines with other arrangement 21

7. Lateral accessory spine on segment 10 Echinoderes dujardinii Claparede, 1863

Lateral accessory spines absent on segments 3-11 8

8. Trunk length greater than 300 /un 9

Trunk length less than 300 Mm 11

9. Lateral spine on segment 12 short (12-17 Mm), less than half the length of lateral spine on

segment 11 Echinoderes pacificus Schmidt, 1974

Lateral spines on segment 12 nearly equal to length of lateral spines on segment 11 10

10. Lateral terminal accessory spines 12%—15% of trunk length, midventral placid wider than

adjacent placids Echinoderes koxloffi, new species

Lateral terminal accessory spines about 8% of trunk length, midventral placid not wider

than adjacent placids Echinoderes pilosus Lang, 1949

11. Middorsal spine on segment 10 slightly longer than middorsal spine on segment 9

Echinoderes ehlersi Zelinka, 1913

Middorsal spine on segment 10 twice as long as middorsal spine on segment 9 12

12. Lateral terminal accessory spines about 41% of trunk length, no lateral accessory spine on

segment 12 Echinoderes worthing Zelinka, 1928

Lateral terminal accessory spines about 23% of trunk length, lateral accessory spine adjacent to lateral spine 12 Echinoderes ferruginous Zelinka, 1928

13. Middorsal spine on segment 11 (juvenile?) Echinoderes sonadiae Timm, 1958

Middorsal spine absent on segment 11 14
4 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

14. Middorsal spine on segment 7 15

Middorsal spine absent on segment 7 18

15. Middorsal spines on segments 6, 7, and 9 only Echinoderes setigera Greeff, 1869

Middorsal spines with other arrangement 16

16. Middorsal spines on segments 6-9 only Echinoderes druxi d'Hondt, 1973

Middorsal spine on segment 5 17

17. Middorsal spines on segments 5-8 only Echinoderes tchefouensis Lou, 1934

Middorsal spines on segments 5-10 only Echinoderes borealis Greeff, 1869

18. Middorsal spines on segments 6 and 9 only Echinoderes citrinus Zelinka, 1928

Middorsal spines on segments 6, 8, and 10 only 19

19. Lateral accessory spines on segments 3, 8-10 with subdorsal spine on segment 3, terminal

tergal extensions evenly tapering to point Echinoderes newcaledoniensis Higgins, 1967

Lateral accessory spine on segment 10 or missing on all segments except terminal segment

of female, terminal tergal extensions with uneven margins 20

20. Lateral accessory spine on segment 10 only, middorsal spines short, (15-18 /*m). Terminal

tergal extensions mesially recurved and somewhat rectangular with small protuberances

at corners Echinoderes riedli Higgins, 1966a

Lateral accessory spines absent on all segments except terminal segment of female, middorsal spines long (73-125 ^m). Terminal tergal extensions rounded with elongate

central spine Echinoderes arlis Higgins, 1966b

21. Posterior border of 12th tergal plate interrupted medially to form broad spine-shaped

process. Small lateral spine or prominent hair (about 8 ^m) on segment 12, similar

spine or hair on segment 13 anterior to lateral terminal accessory spine of female

Echinoderes bookhouti Higgins, 1964b

Posterior border of 12th tergal plate even. Lateral spine or prominent hair absent on

segment 12, segment 13 with lateral terminal accessory spine of female and lateral terminal

spine only 22

22. Lateral spine on segments 7-11 only 23

Lateral spines with other arrangement 25

23. Lateral terminal spines short, about 17% of trunk length. Spinous fringe most prominent

at posteroventral margin of segment 4 and at lateral margins of sternal-tergal junctions

of segments 9 and 10

Echinoderes brevicaudatus, substitute name for Echinoderes brevispinosus Higgins, 1966a

Lateral terminal spines 24%-40% of trunk length, spinous fringe uniform on each segment

where it occurs 24

24. Terminal sternal plates pointed, tergal plate extensions displaced medially from lateral

terminal spines, midventral placid expanded at anterior margin, prominent spinous

fringe along entire posteroventral margin of segment 3 ... Echinoderes remanei (Blake, 1930)

Terminal sternal plates rounded, tergal plate extensions displaced laterally alongside

lateral terminal spines, midventral placid evenly rounded or slightly truncate at anterior

margin, spinous fringe more prominent near midline of posteroventral margin of

segment 3 Echinoderes pennaki Higgins, 1960

25. Lateral spines on segments 6-9 only Echinoderes canariensis Greeff, 1869

Lateral spines with other arrangement 26

26. Lateral spines on all segments (probably error in description)

Echinoderes steineri (Chitwood, 1951)

Lateral spines on segments 8-11 only 27

27. Lateral terminal spines about 50% of trunk length, lateral spine on segment 11 same

length as or slightly longer than that of segment 10

Echinoderes elongata (Nyholm, 1947b)

Lateral terminal spines equal to or longer than trunk length, lateral spine of segment 11

very long Echinoderes levanderi Karling, 1954

Echinoderes dujardinii C llaparede "Echinoceras" Leuckart, 1854:355 [erroneous spelling;

Helgoland].

FIGURES 1-12 Echinoderes Dujardinii Claparede, 1863:90-92, pi. 16: figs.

7-13 [type-locality: St. Vaast la Hougue].—Leuckart, 1869:

"I'Echinodere" Duprdin, 1851:158, pi. 3: figs. 1-4 [St. Malo].— 300.—Greeff, 1869:88-89, pi. 4: figs. 1-5 [by inference:

Schultze, 1853:253. Ostende, Xieuwpoort, Dieppe, Canary Islands; incorrectly
NUMBER 248

cites Dujardin as author].—Panceri, 1876:4, 5, figs. 6, 7

[Ischia, incorrectly cites Greef (sic) as author].—Reinhard,

1881:589 [Odessa, incorrectly cites Dujardin as author].—

Moebius, 1884:7, 1887:117.—Reinhard, 1885:5.—Cams,

1885:184.—Ludwig, 1886:882.—Reinhard, 1887:438 [incorrectly cites Dujardin as author].—Perrier, 1893:453.—

Zelinka, 1896:199 [incorrectly cites Greeff as author].—

Grobben, 1905:336.—Schepotieff, 1907b:297, 298, pi. 17:

figs. 1-6, 16a, pi. 18: figs. 22, 23, pi. 20: fig. 13 [Bergen,

Naples, Brindisi, Rovignj].—Southern, 1914:69, 71 [Clew

Bay, Blacksod Bay].—Zelinka, 1928:221 [lapsus calami}].—

Abe, 1930:39-43.

Echinodera Dujardinii Gosse, 1864:403-404, pi. 2: fig. 16 [decription taken from Dujardin, 1851 only].—Zelinka, 1928:

228 [synonomy].

Not Echinoderes Dujardinii.—Metschnikoff, 1865:459-461

[Helgoland, =Echinoderes subfuscus Zelinka, 1928]; 1869:

190-193 [Selerno, = Echinoderes subfuscus Zelinka, 1928].

Echinoderes brevispinosa Metschnikoff, 1869:190 [Salerno].—

Leuckart, 1869:300-301.—Reinhard, 1887:438.

Echinoderes Sieboldii Pagenstecher, 1875:117-123, pi. 7: figs.

1-6 [Palma]; 1877:88-89, fig. 69.—Cams, 1885:185.—

Reinhard, 1887:438 [Odessa].—Schepotieff, 1907b:298, 299,

pi. 17: fig. 7.—Abe, 1930:39-42.

Echinoderes' brevispinosus.—Panceri, 1876:4 [correction of

E. brevispinosa Metschnikoff, 1869].—Zelinka, 1928:228

[synonomy].

Echinoderes meridionales Panceri, 1876:5, 6, fig. 8 [Ischia].—

Carus, 1885:185.—Zelinka, 1928:254, pi. 2: figs. 11-13

[synonomy, =Habroderes meridionales'].

Echinoderes dujardiniiQ).—Hartog, 1896:236, fig. 120 [Worthing].—Mclntyre, 1962:503.

Echinoderes dujardini.—Hartog, 1896:237.—Remane, 1928:75,

76, figs. 13, 15 [incorrectly cites Greeff as author on p. 76

and in figure legends].—Johnston, 1938:5.—Higgins, 1960:

88-90.—Mclntyre, 1962:503.—Higgins, 1964a: 244, 246;

1964b:489-491; 1966a:120; 1967:75,79; 1969b:113; 1971:25.—

Kozloff, 1972:121.—Moore, 1973:349.—Higgins, 1974:512.—

Boykin, 1974:40.—Schmidt, 1974:12.

Echinoderes dujardinii.—Levander, 1900:20.—Schepotieff,

1907a: 147, 148, figs. 10-12 [Bergen, Naples].—Zelinka, 1907:

132; 1908:630-641, figs. 1, 5, 6, 9 [incorrectly cites Greeff as

author]; 1912:520-527 [incorrectly cites Greeff as author];

1913:419, 420 [incorrectly cites Greeff as author]; 1928:221

[incorrectly cites Greeff as author], 228 [cites Greeff as

author in synonomy], 228-235, figs. 17-19; pi. 3: figs. 1,

2, pi. 10: figs. 1-5, 7-23 [Naples, Trieste, Kiel].—Remane,

1928:60, 62, figs. 7, 9 [incorrectly cites Greeff as author].—

Lou, 1934:3.—Remane, 1936: 333, figs. 215, 252, 266.—

Zaneveld, 1938:261 [Scheveningen].—Nyholm, 1947a:424

[Gullmar Fjord]; 1947b:5, fig. 1.—Lang, 1949:3, 17.—

Tokioka, 1949:67.—Hyman, 1951:180.—Karling, 1954:189-

192.—Kirsteuer, 1964:389—Marinov, 1964:62, 63, fig. 2

[Varna, doubtful record].

Echinoderes sieboldi.—Zelinka, 1928:228 [synonomy].—

Remane, 1936:344 [synonomy].

Habroderes dujardinii Zelinka, 1928:248-250, pi. 4: figs. 1, 2

[described as "n. 1. (zu Echinoderes dujardinii Clap.)"].—

Remane, 1936:332, 333, fig. 273 [noted as larval stage of

Echinoderes dujardinii].

Habroderes meridionales Zelinka, 1928:254, pi. 2: figs. 11-13

[synonomy].—Remane, 1936:333 [noted as larval stage of

Echinoderes dujardinii].

Centropsis parallela Zelinka, 1928:269, pi. 1: figs. 4, 5, 11, 12

[Naples, Trieste].—Remane, 1936:332, 333, fig. 273 [noted

as larval stage of Echinoderes dujardinii].

Echinoderes Masudai Abe, 1930:42, 43, pi. 1: figs. 1, 2 [Hiroshima.].—Remane, 1936:345, fig. 277.—Tokioka, 1949:67.

Echinoderes sieboldii.—Lou, 1934:3 [erroneous spelling].

Echinoderes masudai.—Lang, 1949:17.—Higgins, 1960:89

[nomen dubium]; 1966a:123; 1967:75; 1971:26.

Echinoderes aff. dujardini.—Bacescu et al., 1963:137, 138, figs.

la-c [Agigea, probably not Echinoderes dujardinii].

Not Echinoderes brevispinosus Higgins, 1966a:l 18-121, figs. 1,

2; 1966b:519.—Schmidt, 1974:14. [=Echinoderes brevicaudatus, substitute name].

Not Echinoderes dujardinii.—Chitwood, 1964a:2 [Tomales

Bay, Calif; identity uncertain].

Not Echinoderes dujardini.—Merriman and Corwin, 1973:

227-243, figs. 1-13 [San Juan Island, Washington; =Echinoderes kozloffi, new species].

REDESCRIPTION.—Adults (Figures 1-6), trunk

length, 328-405 um; MSW-9, 78-85 urn, 19.9-24.9

percent of trunk length; SW, 70-83um, 18.9-24.9

percent of trunk length.

Second segment with 16 anteriorly rounded

placids, midventral placid truncate, expanded

slightly at anterior margin, distinctly larger than

adjacent placids; trichoscalid plates on sides of midventral placid with medial indentation on anterior

margin, posterior margin expanded laterally.

Segments 3-12 with short hairs, pattern distinctive (Figures 1, 2); posterior border of segments

5-11 with fine pectinate fringe ventrally, segments

6-11 with pectinate fringe dorsally; terminal segment with few hairlike processes along posterior

margin, posterior margin of tergal plate deeply

incised forming pointed extensions mesial to base

of each lateral terminal spine, sternal plates broadly

rounded with spinous extensions, 10-12 urn in

length (Figures 3, 5).

Middorsal spines on segments 6-10, increasing

uniformly in length, 13-29 um; lateral spines on

segments 4 and 7-12, 14-29 um in length; L-4, 7,

10 and 12 each with adhesive gland at base; L-10

accompanied by dorsally adjacent lateral accessory

spine of nearly equal length, 21-26 ^m; L-4 usually

longer (mean 24.2 um) than remaining lateral

spines; lateral terminal spines long, 160-192 um,

40.7-54.8 percent of trunk length; lateral terminal

accessory spines of female 52-62 um in length; males

without lateral terminal accessory spines but with

penis spines in same position, anteriormost penis
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

K I

100pm

1 2

FIGURES 1-2.—Echinoderes dujardinii, adult female (USNM 53342, RH125.14), neck and trunk

segments: 1, ventral view; 2, dorsal view.
NUMBER 248

50pm

6

FIGURES 3-6.—Echinoderes dujardinii, segments 12-13: 3, ventral view, lateral half, adult female

(USNM 53342, RH125.14); 4, dorsal view, lateral half of same adult female; 5, ventral view,

lateral half, adult male (USNM 53342, RH125.35); 6, dorsal view, lateral half of same

adult male.

spine (P-l) 23-33 um in length, mesially adjacent

penis spine (P-2) 24-36 um, posteriorly adjacent

penis spine (P-3) 26-39 um in length.

Pachycycli (thickened anterior margins of trunk

segments) well developed, forming a distinctive pattern at ventral midline and at attachments of lateral

terminal spine muscles on segment 13; distinctive

muscle scars on ventrolateral portion of first trunk

segment, similar (but reversed orientation) lateral

scars on eighth sternal plates; sensory spots, 2-3 um

in diameter, situated middorsally on segments 3-5,

with two such spots on segment 12 (possibly one on

each of segments 12 and 13), dorsolaterally on segments 6-11, and ventromesially on segments 5-12

(Figures 1, 2).

Morphometric data for adult specimens are

shown in Table 1.

JUVENILE STAGES.—Preadult stage (J-6, "Habroderes-stage," Figures 7, 8) trunk length, 320-328 um;

estimated MSW-9, 78-80 (im, 24.3-25.6 percent of

trunk length; estimated SW, 65-72 um, 19.8-23.5

percent of trunk length (estimated since tergalsternal junctions are not defined in juvenile stages).

Second segment similar to that of adult, both

placids and trichoscalid plates less well developed.

Trunk segments with fewer hairs than adult,

pattern less distinctive and more variable; posterior

borders of segments without pectinate fringe, with

hairs (striations?) along border, group of prominent

hairs (striations?) at ventral midline, terminal segment slightly incised dorsally and ventrally, sternal

area without spinous extensions, tergal area with

small extensions 2-4 um in length, not evenly

tapered.

Middorsal spines on segments 6-11, increasing

uniformly in length, 17-39 um; lateral spines on

segments 4 and 7-12, 17-26 jun in length; L-4, 7,

10 and 12 with poorly developed adhesive glands

at base; L-10 and L-12 accompanied by a dorsally

adjacent lateral accessory spine of nearly equal

length, 21-25 um, but thinner; L-4 more prominent

than remaining lateral spines; lateral terminal

spines long, 148-152 um, 45.1-48.7 percent of trunk

length; lateral terminal accessory spines 42-47 ^m

in length, sexes often indistinguishable in juvenile

stages unless developing oocytes visible.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 1.—Measurements (/im) and indices (%) for Echinoderes dujardinii adults

TL

SU

SW/TL

HSW-9

MSH/TL

Om

Dm/TL

D-6

0-7

D-8

D-9

D-10

Lm

Lm/TL

a ...

at...

<t ...

9 .. .

<t9 ...

<f ...

9 .. .

d"9 .. .

<s ...

9 ...

d9 ...

a

9 ...

d9 .. .

<f .. .

9 .. .

rf9 .. .

<f .. .

9 .. .

<fi ...

a ...

9 .. .

tf9 ...

<f ...

9 ...

* 9 .. .

cf .. .

9 .. .

cf9 .. .

cf .. .

9 ...

<*9 .. .

tf .. .

9 .. .

<*9 .. .

9 ...

•*•«.. .

C .. .

9 ...

<f9 ...

26

20

46

26

19

45

26

19

45

26

19

45

26

19

45

10

17

27

10

17

27

11

19

30

18

17

35

18

19

37

20

19

39

23

20

43

18

17

35

18

17

35

328-380

336-405

328-405

70-82

75-83

70-83

19.3-24.9

18.9-24.4

18.9-24.9

75-85

78-85

75-85

20.7-24.9

19.9-24.7

19.9-24.9

15.8-18.8

15.0-19.8

15.0-19.8

4.6-5.5

4.0-5.6

4.0-5.6

13-16

13-16

13-16

13-19

13-18

13-19

11-23

14-21

11-23

13-23

16-23

13-23

20-26

16-27

16-27

17.7-22.8

19.2-25.0

17.7-25.0

4.6-6.7

4.9-6.9

4.6-6.9

350.5

367.9

358.0

76.6

78.9

77.0

21.6

21.5

21.5

79.4

81.5

80.2

22.7

22.1

22.4

17.3

17.6

17.5

5.0

4.8

4.8

14.0

14.2

14.1

15.3

15.5

15.4

16.3

17.1

16.7

18.6

19.5

19.0

22.6

21.9

22.3

20.6

21.2

20.9

5.8

5.8

5.8

Standard

15.4

21.9

18.3

2.7

2.6

2.6

1.3

1.5

1.4

2.6

1.8

2.3

1.3

1.5

1.4

1.1

1.7

1.4

3.2

0.4

0.1

1.0

1.1

1.1

2.0

1.4

1.7

1.1

1.7

1.4

2.6

2.7

2.6

2.4

3.1

2.7

1.4

1.5

1.4

0.5

0.1

0.5

Standard

3.0

4.9

2.7

0.5

0.6

2.6

0.2

0.4

0.2

0.5

0.4

0.3

0.2

0.3

0.2

0.4

0.4

0.3

0.1

0.1

0.02

0.3

0.3

0.2

0.5

0.3

0.3

0.3

0.4

0.2

0.6

0.6

0.4

0.5

0.7

0.4

0.3

0.4

0.2

0.1

0.6

0.1

Coefficient

4.3

6.0

5.1

3.6

3.3

3.4

5.8

7.1

6.3

3.3

2.2

2.8

5.5

6.9

6.1

6.3

9.4

8.3

6.3

9.7

2.6

7.1

8.0

7.6

12.9

8.8

10.9

7.0

9.7

8.9

13.9

14.0

13.7

10.4

14.0

12.0

6.9

6.8

6.8

8.5

9.9

9.0

L-4 (AT)

L-7

L-8

L-9

L-10

LA-10

L-ll

L-12

LTS

LTS/TL

LTAS

LTAS/TL

P-l

P-2

P-3

9 ...

cT9 .. .

a

9 ...

tf9 ..

cf ..

9 ..

cf9 ..

a ..

9 ..

<**

<f ..

9 ..

cf9 ..

a ..

9 ..

rf9 ..

d ..

9 ..

cf9 ••

a ..

9 ..

cf9 ..

a ..

9 ..

tf9 ..

cf

9 ..

<*9 ..

9 ..

9 ..

a ..

<s ..

a

16

17

33

19

17

36

26

19

45

26

19

45

26

20

46

22

18

40

25

20

45

26

19

45

26

20

46

26

20

46

T9

19

25

26

19

17-26

22-28

17-28

13-25

14-26

13-26

14-22

16-26

14-26

17-23

16-28

16-28

20-25

18-26

18-26

22-26

21-26

21-26

17-28

20-27

17-28

20-29

20-28

20-29

160-184

160-192

160-192

44.3-54.8

40.7-54.0

40.7-54.8

52-62

13.7-17.5

23-33

24-36

26-39

23.0

25.2

24.2

17.8

18.9

18.3

18.0

19.3

18.6

20.4

20.6

20.5

22.2

22.2

22.2

24.6

24.1

24.4

22.5

22.9

22.7

23.0

24.4

23.6

175.5

176.6

175.6

50.2

47.9

49.2

56.6

15.3

27.4

30.6

33.5

Standard

2.3

1.6

1.9

3.6

3.5

3.5

2.3

2.7

2.4

2.1

2.8

2.4

1.2

1.6

1.4

1.5

1.6

1.5

2.1

1.8

2.0

1.9

2.1

1.9

6.1

8.6

7.2

2.9

3.8

3.0

3.1

1.2

2.9

3.1

3.9

Standard

0.6

0.4

0.3

0.8

0.8

0.6

0.4

0.6

0.4

0.4

0.6

0.4

0.2

0.4

0.2

0.3

0.4

0.2

0.4

0.4

0.3

0.4

0.5

0.3

1.2

1.9

1.1

0.6

0.8

0.5

0.7

0.3

0.6

0.6

0 . 9

Coefficient

9.8

6.2

7.9

20.2

18.3

19.0

12.6

13.9

13.1

10.5

13.6

11.8

5.6

7.4

6.3

6.2

6.5

6 . 3

9.5

7.8

8.6

8.2

8.5

8.2

3.5

4.9

4.1

5.7

7.8

6.6

5.4

7.8

10.5

10.0

11.5

Pachycydi not well defined, muscle scars on ventrolateral areas of first trunk segment reduced, no

scars on segment 8; sensory spots, 2-3 um in diameter, situated middorsally on segments 4-5 and

12-13, laterodorsally on segments 6-13, mesially on

sternal plates 4-11.

Fourth-stage juvenile (J-4, "Habroderes-stage,"

Figures 9, 10) trunk length, 224 jun; estimated

MSW-9, 72 urn, 32 percent of trunk length; estimated SW, 65 um, 29 percent of trunk length.

Second segment as in preadult, less well developed.

Trunk segments similar to those of preadult but

not well defined posteriorly so that only 10 trunk

segments apparent; series of medial hairs (striations?) both dorsally and ventrally indicating area

of presumptive segment 12; corresponding area of

segment 13 less distinct; hairs less obvious than in

preadult, more scattered; terminal segment slightly

incised ventrally with suggestions of pointed tergal

extensions.

Middorsal spines on segments 6-11, D-6-10,

14-39 \tsn, increasing uniformly in length to segment 11, D-ll, 79 um, twice the length of D-10,

extending slightly beyond terminal segment; lateral

spines similar to preadult stage, 16-26 um in length;

lateral terminal spines 160 um in length, 71.4 percent of trunk length; lateral terminal accessory

spines 47 um.

Third-stage juvenile (J-3, "Hapaloderes-stage,"

Figures 11, 12) trunk length, 200-208 um; estimated

MSW-9, 72 um, 34.3 percent of trunk length; estimated SW, 51 (wn, 24.5 percent of trunk length.

Second segment as in J-4 stage, less well developed.

Trunk segments similar to those of J-4 stage, less
NUMBER 248

,V (A.

\ . ' 1

100pm

, -J _

fj}ii MA

FIGURES 7-8.—Echinoderes dujardinii, preadult (J-6) stage (USNM 53345, RH125.49), neck and

trunk segments: 7, ventral view; 8, dorsal view.
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

100pm

FIGURES 9-10.—Echinoderes dujardinii, juvenile J-4) stage (USNM 53344, RH125.53), neck and

trunk segments: 9, ventral view; 10, dorsal view.
NUMBER 248 11

100pm

I , f

I I

FIGURES 11-12.—Echinoderes dujardinii, juvenile (J-3) stage (USNM 53343, RH125.54), neck and

trunk segments: 11, ventral view; 12, dorsal view.

well defined posteriorly, striations along posterior tions incomplete on presumptive segment 10, remargins of segments 3-9 defining segments, stria- stricted to midventral area, striations restricted to
12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

middorsal and midventral areas of presumptive segment 11, striations apparent on midventral area of

presumptive segment 12; terminal segment blunt,

without extensions.

Middorsal spines on segments 6-12 (D-12 essentially midterminal), D-6-9, 17-29 um, increasing

uniformly in length to segment 10, D-10, 39-43

urn in length, D-ll, 98-109 urn in length, D-12

(midterminal) 136-170 um in length, 60.7-85.0 percent of trunk length; lateral spines similar to J-4

stage except LA-12 absent, 18-22 um in length;

lateral terminal spines 60-64 \im, 28.8-30.0 percent

of trunk length; lateral terminal accessory spines

46 um.

MATERIAL EXAMINED.—46 adults (USNM 53342) consisting

of 26 males and 20 females, 4 preadults (J-6, "Hapaloderesstage") (USNM 53345), one fourth-stage juvenile (J-4, "Hapaloderes-stage") (USNM 53344), and 2 third-stage juveniles

(J-3, "Habroderes-suge") (USNM 53343); col. Dr. Eugene N.

Kozloff, 19 October 1973, Roscoff, Nord-Finistere, France

(from the surface of sandy mud in the port).

REMARKS.—Dujardin (1851) and many subsequent authors prior to Zelinka (1928) introduced

certain ambiguities into the description of Echinoderes dujardinii. The most common problems include the interpretation of the animal's length and

the position of the spines on the trunk segments.

Dujardin, for example, probably included the head

and neck and may have included some juvenile

stages in his total length measurements (300-550

um). Prior to Zelinka (1928), most authors were not

consistent in numbering segments; consequently,

one must carefully compare the illustrations with

the text when assessing the correct position of middorsal and lateral spines.

The most accurate illustration of E. dujardinii

published prior to Zelinka (1928) was that of Greeff

(1869). One of the most important characteristics of

this species, the presence of two lateral spines on

segment 10, was noted by Dujardin (1851), but

ignored by many subsequent authors.

Including E. dujardinii, the adults of seven species of Echinoderes have middorsal spines on segments 6-10 and lateral spines on segments 4, 7-12

(£. pilosus Lang, 1949 may have lateral spines or

adhesive tubes on segment 4 although they are not

included in the author's description). Echinoderes

ehlersi Zelinka, 1913 (from Zanzibar) is 228 um

long, much smaller than E. dujardinii, and has significantly longer lateral terminal spines in relation

to the trunk length, 77 percent as contrasted with

41-55 percent in E. dujardinii. Echinoderes ferrugineus Zelinka, 1928 has been found with E. dujardinii at both Naples and Trieste (Zelinka, 1928).

Echinoderes ferrugineus also is smaller than E. dujardinii and has longer lateral terminal spines

relative to its trunk length, about 67-77 percent.

Echinoderes ferrugineus differs from E. ehlersi

in that D-10 in the former species is 55 urn, nearly

twice the length of D-9; in E. ehlersi, the middorsal spines increase uniformly from 8-14 um in

length.

Echinoderes worthingi Zelinka, 1928 also occurs

with E. dujardinii. One specimen was collected

along with E. dujardinii in this study and Southern

(1914) found both species at Blacksod Bay, Ireland.

Echinoderes worthingi is similar in size and other

characteristics to both E. ehlersi and E. ferrugineus

as noted by Zelinka (1928). Echinoderes worthingi

most closely resembles E. ferrugineus in that the

length of D-10, 45-50 um, is twice that of D-9,

19-23 um. The two species appear to be distinguishable by the shape of the terminal segment, a slightly

smaller lateral terminal spine, and the presence of

a prominent sensory hair (spine?) posteriorly adjacent to the lateral spine on segment 12 in E. ferrugineus; a similar hair exists in E. pacificus

(Figures 28, 30).

Three additional species with the D-6-10, L-A,

7-12 spine combination are also similar to E.

dujardinii in size. Echinoderes pilosus (from South

Georgia Island) is poorly described, and as mentioned previously, I am including it because there

is a chance that Lang (1949) overlooked the lateral

spines (= adhesive tubes) on segment 4. Echinoderes pilosus differs from E. pacificus Schmidt,

1974 (from the Galapagos) in that the former has

relatively longer lateral terminal spines, about 52

percent of the trunk length compared with 27-36

percent in the latter. If Lang is correct, a distinguishing feature of E. pilosus is the uniform size of

the placids. In the other species, however, the midventral placid is much wider than the adjacent

ones. Both E. pacificus and the remaining species,

E. kozloffi, new species, are discussed more extensively below.

All six species mentioned in the foregoing discussion differ from E. dujardinii by their lack of a

lateral accessory spine on segment 10.

Two additional species of Echinoderes share with
NUMBER 248 13

E. dujardinii the presence of a lateral accessory

spine on segment 10. These include E. riedli Higgins, 1966a (from the Red Sea) and E. newcaledoniensis Higgins, 1967. Echinoderes riedli is

smaller, 238 \un in length. Echinoderes newcaledoniensis is the only member of this genus having

lateral accessory spines on segments 8-11 and is

unique in possessing dorsolateral spines on segment

4. Both E. riedli and E. newcaledoniensis differ

from E. dujardinii by having middorsal spines on

segments 6, 8, and 10 only.

DISTRIBUTION.—Echinoderes dujardinii has been

reported from 26 localities (Figure 13, also see annotations in synonomy). Most of the reports are

European and include both the northern and

southern coastal areas of the continent. These distribution records are probably reliable except for

those from the Black Sea. Based on the evidence

presented by Reinhard (1881), Bacescu et al. (1963),

and Marinov (1964), I believe these records are

questionable. Similarly, the reports of E. dujardinii

from Japan (Tokioka, 1949) and the Pacific coast

of the United States (Chitwood, 1964a; Merriman

and Corwin, 1972) are based on misidentifications

of E. kozloffi, new species.

Echinoderes kozloffi, new species

FIGURES 14-21

Echinoderes.—Kozloff, 1972:121, figs. 1-18—Higgins, 1974:514,

figs. 11-16.

Echinoderes dujardini.—Merriman and Corwin, 1973:227-243,

figs. 1-13.

Echinoderes sp.—Boykin, 1974:40.

DIAGNOSIS.—Trunk length, 328-376 jim, middorsal spines on segments 6-10, increasing uniformly in length; lateral spines on segments 4, 7-12,

with adhesive glands at base of L-4, 7, 10 and 12,

lateral spine of segment 12, 20-30 um in length;

lateral extensions of terminal tergal plate tapering

evenly to point protruding about 5 um beyond

extensions of tergal plate (exclusive of prominent

spines on latter).

DESCRIPTION.—Adults (Figures 14-19), trunk

length, 328-376 um; MSW-9, 68-74 um, 18.0-22.3

percent of trunk length; SW, 62-69 urn, 17.5-20.0

percent of trunk length.

Second segment with 16 anteriorly rounded placids; midventral placid truncate, not expanded laterally at anterior margin, distinctly larger than

adjacent placids; trichoscalid plates on sides of

FIGURE 13.—Echinoderes dujardinii, distribution records, open circles indicate doubtful records.
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

'/r^/

""»«*,.

^iffi

te'ito

wu

100pm

14 15

FIGURES 14-15.—Echinoderes kozloffi, new species, holotypic female (USNM 5SS37), neck and

trunk segments: 14, ventral view; 15, dorsal view.
NUMBER 248 15

midventral placid indented at anteromedial margin,

posterior margin expanded laterally, four trichoscalid plates on dorsal placids much smaller than

those on ventral placids.

Segments 3-12 with short hairs, pattern distinctive (Figures 14, 15); posterior border of segments

4-11 with fine pectinate fringe dorsally and ventrally; terminal segment with few hairlike processes

along posterior margin; posterior margin of tergal

plate deeply incised, laterally forming pointed,

evenly tapered extensions mesial to base of each

lateral terminal spine; sternal plates broadly

rounded with spinous extensions, 15-22 um in

length (Figures 16, 18).

Middorsal spines on segments 6-10, increasing

uniformly in length posteriorly, 20-45 um; lateral

spines on segments 4 and 7-12, 24-32 um in length;

L-4, 7, 10 and 12 with adhesive gland at base; L-8

usually shorter (mean 22.9 um) than L-7 (mean 27.5

um) or L-9 (mean 26.1 um); L-9-12 all similar in

length, 22-32 um; lateral terminal spines long, 144-

180 um, 42.0-52.4 percent of trunk length; lateral

terminal accessory spines of female 46-59 um in

length; males without lateral terminal accessory

spines but with penis spines in same position;

anteriormost penis spine (P-l) 25-33 um in length,

mesially adjacent penis spine (P-2) 21-27 um in

length and distinctly broader, posteriorly adjacent

penis spine (P-3) 35-42 um in length, generally

tending to curve mesially, crossing lateral terminal

spine.

Pachycycli well developed, forming distinctive

pattern at ventral midline and superficial to lateral

terminal spine muscle attachments on segment 13;

muscle scars on ventrolateral portion of first trunk

segment almost indistinguishable, similar V-shaped

scars on either side of ventral midline of segments

4-9; prominent scars on lateral margins of sternal

plates 8-9, more centrally located on sternal plates

of segment 10 (Figure 14); sensory spots, 2-3 um in

diameter, situated middorsally on segments 3—5

with two such spots on segment 12 (possibly one on

each of segments 12 and 13), dorsolaterally on seg50pm

18 19

FIGURES 16-19.—Echinoderes kozloffi, new species, segments 12-13: 16, ventral view, lateral half,

holotypic female (USNM 53337); 17, dorsal view, lateral half, holotypic female; 18, ventral

view, lateral half, allotypic male (USNM 53338); 19, dorsal view, lateral half, allotypic male.
16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ments 3-11, ventromesially on segments 4-12, anterolaterally on ventral surface of segment 3, and

possibly near lateral margin of sternal plates 5-7

(Figures 14, 15).

Morphometric data for adult specimens are

shown in Table 2.

JUVENILE STAGES.—Preadult stage (J-6, "Habroderes-stage," Figures 20, 21) trunk length, 276-

300 um; estimated MSW-9, 64-68 um, 23-24 percent of trunk length; estimated SW, 62-64 um,

22-23 percent of trunk length (estimated since

tergal-sternal junctions not defined in juvenile

stages).

Second segment similar to that of adult, both

placids and trichoscalid plates less well developed.

Trunk segments with fewer hairs than adult although hairs as long as in adult (5-9 um), pattern

less distinctive and more variable; posterior borders

of segments without pectinate fringe, with hairs

(striations?) along border, group of prominent hairs

(striations?) at ventral midline and second group

lateral to midline; terminal segment slightly incised

dorsally and ventrally, sternal area without spinous

extensions, tergal area with evenly tapered extensions, 2-4 um in length.

Middorsal spines on segments 6-11, increasing

uniformly in length, 17-62 um (mean 27.9-34.0 urn,

10.7-12.0 percent of trunk length), D-ll extending

beyond terminal tergal borders; lateral spines on

segments 4, 7-12, 17-26 um in length; L-4, 7, 10,

and 12 with poorly developed adhesive glands at

base; L-12 accompanied by a lateral accessory spine

situated dorsally to it, slightly longer (22-26 um)

and thinner; L-4 also thinner than other lateral

spines; lateral terminal spines long, 152-160 um,

50.7-56.5 percent of trunk length; lateral terminal

accessory spines 35-40 um in length.

Pachycycli not well defined, muscle scars similar

to those of preadult stage of E. dujardinii.

Fifth-stage juvenile (J-5, "Habroderes-stage")

trunk length, 224-256 um; other measurements similar to preadult stage described above.

TABLE 2.—Measurements Qua) and indices (%) for Echinoderes kozloffi, new species, adults

Char

Tl

SW

SW/TL

MSW-9

MSW/TL

Di

Dn/TL

D-6

D-7

D-8

D-9

D-ia

I n

acter

a .

9 .

rf«.

a .

9 .

d9.

<f .

9 .

fi.

tt .

9 .

ri.

tt .

9 .

cf9.

tf .

9 .

c«.

a .

9 .

rfV.

a .

9 .

a 9.

a .

9 .

d 9 .

a .

9 •

tf9.

d" .

9 .

cf9.

cf .

9 .

O9.

rf .

9 .

O9.

Number

20

20

40

19

20

39

19

20

39

19

20

39

19

20

39

9

11

20

9

11

20

10

14

24

13

12

25

13

15

28

14

16

30

20

20

40

20

20

40

Range

328-364

332-376

328-376

62-66

62-69

62-69

17.7-19.8

17.5-20.0

17.5-20.0

68-72

69-74

68-74

18.8-21.4

18.0-22.3

18.0-22.3

26.6-29.8

25.4-29.2

25.4-29.8

7.6-9.1

7.1-8.1

7.1-9.1

20-26

20-26

20-26

21-27

22-26

21-27

23-:?

22-26

22-27

26-30

24-33

24-33

38-45

36-43

36-15

24.7-27.7

24.0-27.7

24.0-27.7

Mean

334.4

356.0

350.2

62.3

66.4

65.4

18.5

18.6

18.6

69.5

71.8

70.7

19.9

20.1

20.1

28.7

27.2

27.9

8.3

7.6

7.9

23.3

22.0

22.5

24.4

23.4

23.9

25.7

24.4

25.0

27.6

26.9

27.2

41.6

39.6

40.6

26.0

26.4

26.2

Standard

Deviation

8.2

12.9

12.2

1.2

1.9

1.9

0.5

0.8

0.7

1.3

1.8

1.9

0.8

1.0

0.8

1.1

1.1

1 .3

0.5

0.3

0.5

2.0

1.9

2.0

1 .8

1.2

1.6

1.0

1.7

1.5

1.3

2.2

1.8

2.5

1.8

2.4

0.8

0.9

0.9

Standard

Error

1.8

2.9

1.9

0.3

0.4

0.3

0.1

0.2

0.1

0.3

0.4

0.3

0.2

0.2

0.1

0.4

0.3

0.3

0.2

0.1

0.1

0.6

0.5

0.4

0.5

0.3

0.3

0.3

0.4

0.3

0.4

0.5

0.3

0.6

0.1

0.4

0.2

0.2

0.1

Coefficient

of Variability

2.4

3.6

3.5

1.8

2.9

2.9

2.5

4.4

3.6

l.S

2.5

2.

3.J

5.1

4.;

4.

4.(

5

4.

6.

8.1

8.

8.

5'

6.

4.0

6.8

6.0

4.9

8.1

6.7

6.1

4.6

5.9

2.9

3.5

3.3

Character

Lm/TL tt .

9 .

O9.

L-4 (AT)d- .

9 .

rf9.

L-7

1-8

L-9

L-10

L-l l

L-12

LTS

LTS/TL

LTAS

LTAS/Tl

P-l

P-2

P-3

a

9

a . ,

9 . .

tf9..

a ..

9 . .

<&..

<f ..

9 . .

09..

a ..

9 . .

<f9..

a ..

9 . .

09..

a ..

9 ••

O9-.

0 ..

9 ••

O9.

9 ... .

V

<f

a

a

Number

20

20

40

20

20

40

20

20

40

20

20

40

20

20

40

20

20

40

20

20

40

20

20

40

20

20

40

20

20

40

20

20

19

20

18

Range

7.1-7.9

6.7-8.1

6.7-8.1

26-29

25-29

25-29

24-29

24-31

24-31

21-25

21-26

21-26

24-28

23-29

23-29

26-30

26-29

26-30

23-30

25-32

23-32

22-29

23-32

22-32

144-172

160-180

144-180

40.0-50.0

43.5-52.4

42.0-52.4

46-59

12.6-15.8

25-33

21-27

35-42

Mean

7.6

7.4

7.5

27.0

27.1

27.0

27.2

27.8

27.5

22.9

22.8

22.9

26.2

26.0

26.1

27.7

27.5

27.6

26.6

27.8

27.2

25.6

26.5

26.0

157.8

170.1

164.0

45.9

47.9

46.9

52.6

14.8

28.2

22.8

38.3

Standard

Deviation

0

0

0

1

1

1

1

1

1

1

1

1

0

1

1

1

1

1

1

1

1

1

2

2

7

6

9

2

2

2

2

4

3

1

3

2

5

6

6

4

3

3

9

3

1

4

2

3

7

7

8

9

4

2

1

7

2

5

5

7

3.5

1.0

2.2

1.8

1.9

Standard

Error

0.1

0.1

0.1

0.2

0.3

0.2

0.3

0.4

0.3

0.3

0.3

0.2

0.2

0.3

0.2

0.3

0.3

0.2

0.4

0.4

0.3

0.4

0.6

0.4

1.6

1.5

1.5

0.6

0.6

0.4

0.8

0.2

0.5

0.4

0.4

Coefficient

of Variability

3.5

5.6

4.5

3.9

4.9

4.4

5.5

5.8

5.7

6.2

5.8

5.8

3.4

5.0

4.2

4.9

4.3

4.6

6.4

6.3

6.7

7.5

9.2

8.5

4.5

4.0

5.6

5.5

5.2

5.7

6.7

6.7

8.0

7.9

4.9
NUMBER 248 17

100pm

20 21

FIGURES 20-21.—Echinoderes kozloffi, new species, preadult (J-6) stage (USNM 53341, RH145.43),

neck and trunk segments: 20, ventral view; 21, dorsal view.
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Middorsal spines on segments 6-11, increasing

uniformly in length, 15-72 um (mean 33.0-39.8

um), 14.7-15.8 percent of trunk length. Note: This

latter statistic distinguishes J-5 from J-6.

HOLOTYPE.—Adult female, TL 344 urn; North

Bay, San Juan Island, Washington, USA (48°31.0'N,

123°O1.O/W); 9 August 1975; col. E. N. Kozloff;

USNM 53337.

ALLOTYPE.—Adult male, TL 348 um; as holotype;

USNM 53338.

PARATYPES.—19 females and 19 males, TL 328-

376 um; data as for holotype; USNM 53339; 5 preadult juveniles, TL 224-300 um, 5 J-5? juveniles,

224-256 um; as holotype; USNM 53340; 4 females

and 4 males, TL 328-364 um; Reid Harbor, Stuart

Island, Washington, USA (48°40.0/N, 123° 1 LOW);

31 July 1963; col. J. C. Boykin; USNM 53347; 3

preadult juveniles, TL 280-292 um; data as for

above; USNM 53348.

REMARKS.—Echinoderes kozloffi, new species,

most closely resembles E. pacificus from the Galapagos Islands. I consider the two sibling species.

A similar sibling relationship involving the two

geographic areas has been noted for two species of

otoplanid turbellarians: Philosyrtis sanjuanensis

Ax and Ax, 1967 (from San Juan Island, Washington) is a sibling species of P. santacruzensis Ax and

Ax, 1974 (from the Galapagos Islands) according to

these authors. Echinoderes kozloffi is similar to E.

pacificus in total length, standard width, maximum

sternal width, and general appearance. The range

of the mean lengths of middorsal spines in E. kozloffi (25.4-29.8 um) and those of E. pacificus (46.5-

56.5 yon) is one of the significant differences between the two species. Other differences include the

longer (25-29 um) lateral spine on segment 4 in

E. kozloffi, which contrasts with the shorter (12-17

um) lateral spine on segment 4 in E. pacificus; in

the former species the lateral terminal spines, 144-

180 um, 42.0-52.4 percent of the trunk length, are

longer than those of the latter species, 90-118 um,

27.0-36.0 percent of the trunk length. The lateral

terminal accessory spines of E. kozloffi are slightly

longer (46-59 um) than those of E. pacificus (34-

41 um).

The relatively long (22-30 um) lateral spine on

segment 12 of E. kozloffi is a particularly noticeable feature that contrasts with the short (12-17

um), blunt lateral spine on segment 12 of E. pacificus. The posterior margins of both tergal and

sternal plates of the terminal segment distinguish

the two species. The mesial margins of the terminal

sternal extensions of E. pacificus are interrupted and

beset with a series of prominent hairs (Figure 28);

in E. kozloffi these margins are even and have a less

obvious series of hairs. Both species have hairs

along the mesial margins of the terminal sternal

extensions but, again, those of E. pacificus are more

distinct, each hair associated with a steplike interruption of the margin. Also, the sternal extensions

of both species have a spinous process. The processes in E. pacificus, however, are half the length

(8-12 um) of those in E. kozloffi (17-25 um). Echinoderes pacificus is discussed later in this paper.

Echinoderes kozloffi is similar to E. pilosus. As

noted above, E. pilosus is not well described but

has the same general trunk length and spination

formula as both E. kozloffi and E. pacificus.

Echinoderes pilosus appears to differ from E. kozloffi in only two principal characters: E. pilosus

has shorter lateral terminal accessory spines, about

35 um, about 8 percent of the trunk length, and

the midventral placid is the same width as adjacent

placids. This latter character is unique within the

genus.

Echinoderes kozloffi is sympatric with E. pennaki

Higgins, 1960 but differs in that the latter species

lacks a lateral spine on segment 12, has conspicuously longer lateral spines (mean 58.1 um, 14

percent of the trunk length), a prominent ventral

spinous fringe on the posterior border of the first

trunk segment, lacks spinous processes on the

terminal sternal plates, and has slightly longer,

more pointed terminal tergal extensions. Other

differences are mentioned in the discussion of

E. pennaki that follows.

ETYMOLOGY.—This species is named in honor of

Eugene N. Kozloff, a fellow student of the

Kinorhyncha.

Echinoderes pennaki Higgins

FIGURES 22-25

Echinoderes pennaki Higgins, 1960:86; 1961:81.—Chitwood,

1964b:3.—Higgins, 1964a:246; 1964b:479—Kirsteuer, 1964:

389.—Higgins, 1966a: 120; 1966b:519; 1967:75.—Schmidt,

1974:13.—Boykin, 1974:40.

REDESCRIPTION.—Holotypic adult female (Figures

22-25), trunk length, 404 um; MSW-8, 62 um, 15
NUMBER 248 19

100pm

FIGURES 22-23.—Echinoderes pennakt, holotypic female (USNM 29746), neck and trunk segments;

22, ventral view; 23, dorsal view.
20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

percent of trunk length; SW, 56 um, 14 percent of

trunk length.

Second segment with 16 anteriorly rounded

placids; midventral placid slightly truncate, distinctly larger than adjacent placids; trichoscalid

plates on sides of midventral placid indented

slightly on anterior margin, short and broad,

trichoscalid plates on dorsal surface similar but

equally as broad as long.

Segments 3-12 with prominent hairs (10-15 um),

pattern distinctive (Figures 22-23); posterior border

of segments 3-11 with pectinate fringe both dorsally and ventrally, fringe on ventral surface of

segment 3 almost spinose, more finely pectinate on

remaining segments, slightly evident near posterior

border of 12th sternal plates; terminal segment

with minute hairs on border of tergal and sternal

plates, posterior margin of tergal plate deeply

incised forming pointed extensions mesial to base of

each lateral terminal spine, sternal plates broadly

rounded without spinous extensions (Figures

24, 25).

50pm

24

FIGURES 24-25.—Echinoderes pennaki, segments 12-13 of holotypic female (USNM 29746): 24, ventral view, lateral half; 25,

dorsal view, lateral half.

Middorsal spines on segments 6-10, increasing

uniformly in length, 48-70 um, mean length 59.6

um, 15 percent of trunk length; lateral spines on

segments 7-11, L-4 (=adhesive tube) absent although a scar, or perhaps a sensory spot, located

slightly mesial to usual position of L-4; L-7

shorter (14 um) than remaining lateral spines (32-

36 um) subequal in length, mean length of lateral

spines 29.8 um, 7 percent of trunk length; lateral terminal spines long, 156 um, 39 percent of

trunk length; lateral terminal accessory spines 52

um in length.

Pachycycli well developed, forming distinctive

pattern at ventral midline and superficial to attachment of lateral terminal spine muscles on segment

13; small muscle scars visible near midventral line

of segments 3-4, 7, 11, and 12, more prominent scars

dorsolaterally on segments 3, 5-12; sensory spots situated ventrolaterally on segment 3 and possibly on

segment 4 (possibly adhesive tube openings?).

MATERIAL EXAMINED.—4 adult females including holotype

(USNM 29746) and 3 paratypes (USNM 29747); col. P. L. Illg,

16 July 1958, East Sound, Orcas Island, Washington, USA,

from a depth of 32 meters.

REMARKS.—Eighteen years of research on the

Kinorhyncha has prompted me to reexamine my

first species descriptions with better experience to

guide me. The combination of this experience and

better optical instruments has allowed me to illustrate and describe E. pennaki more accurately. For

example, the range of "total length" given in my

original description (Higgins, 1960) is "390-430 um

(taken along dorsal surface between anterior edge

of second zonite and posterior edge of zonite 13)."

More standard measurements are now taken from

the anterior edge of segment 3 to the posterior edge

of segment 13. Similarly, the maximum width of

80-90 um originally given for this species differs

from the more precise and standardized measurement of maximum sternal width as noted in previous publications (Higgins, 1967).

Echinoderes pennaki most closely resembles E.

remanei (Blake, 1930) redescribed by me (Higgins,

1964a). The two species differ in size: E. pennaki is

larger (380-404 um) than E. remanei 282-358 um)

yet both have the same spination formula (D-6-10,

L-4? 7-11). Both have a prominent spinous fringe

on the posteroventral border of segment 3; this

feature is more prominent in E. remanei and also

occurs on the fourth segment. The midventral
NUMBER 248 21

placid of E. remanei is expanded laterally along its

anterior margin, and the border of the terminal

sternal plates is pointed, not rounded as in E. pennaki (Figure 24). Our understanding of both species

would benefit from an expanded study of their

taxonomic characters based on larger numbers of

individuals in a given sample.

Echinoderes pacificus Schmidt

FIGURES 26-31

Echinoderes pacificus Schmidt, 1974:1.

This species has been described with considerable

accuracy by my colleague, Dr. Peter Schmidt (1974).

In studying the holotype and paratype material

sent to me by him, however, certain additional

information was revealed by mounting specimens

in Hoyer's and observing them with differential

interference contrast optics. I have reillustrated

E. pacificus (Figures 26-31) in order to facilitate

the comparison of this species with others that

might be confused with it.

Segments 3-12 are covered with short hairs in a

distinctive pattern (Figures 26, 27). A fine pectinate

fringe is evident ventrally on the posterior border

of segments 4-13 although it is much less distinct

on segment 12; segments 6-13 exhibit this fringe

dorsally.

As noted in the discussion of E. kozloffi, the

short (12-17 urn) lateral spine on segment 12 of

E. pacificus is diagnostic. This spine tends to curve

away from the trunk. Slightly posterior to it, at the

junction of segments 12 and 13, there is a sensory

hair (Figures 28-31) not reported in the original

description.

Small muscle scars (2-3 \im in diameter), similar

to sensory spots, are present anterolaterally on the

ventral surface of the first trunk segment. Narrow,

slitlike scars occur posterior to the sensory spots

on sternal plates 7-9. More distinctive muscle

scars (Figure 26) occur near the lateral margin of

the sternal plates of segment 9; similar scars are

situated near the sensory spots of the 10th sternal

plates. Sensory spots occur on either side of the

ventral midline of segments 4-12. Three spots are

present on the dorsal surface of the first trunk

segment; only the median spot persists on the following two trunk segments (segments 4-5). Sensory

spots on the segments bearing middorsal spines

are on both sides of the midline. Two middorsal

sensory spots appear to be on segment 12; one of

these may be associated with segment 13.

MATERIAL EXAMINED.—Holotypic male, TL 372 Mm (USNM

53335); 8 paratypic females and 7 paratypic males (USNM

53336); col. P. Schmidt, July-nSeptember 1972, Academy Bay,

Station IX, 5c, upper subtidal sediments.

REMARKS.—As noted in the discussion of E. kozloffi, this species closely resembles E. pacificus. Both

species share the same spination formula with

E. pilosus, E. ehlersi, E. worthingi, and E. ferrungineus (E. dujardinii is similar but has a lateral

accessory spine on segment 10).

Echinoderes ehlersi, E. worthingi, and E. ferrungineus are smaller, 210-260 \ua trunk length,

although the latter species shares with E. pacificus

the presence of a sensory hair posterior to the

lateral spine on segment 12. Echinoderes pacificus

is most easily distinguished from all species having

the same spination formula by its short lateral spine

on segment 12 and the prominent border of hairs

on the terminal sternal plates.

Echinoderes brevicaudatus, substitute name

for £. brevispinosus Higgins

During the course of the present investigation,

I discovered that the name E. brevispinosa was first

used by Metschnikoff (1869:190), corrected to E.

brevispinosus by Panceri (1876:4), and synonomized

with E. dujardinii by Zelinka (1928:228). Since I

inadvertently applied this preoccupied name to a

species from the Red Sea (Higgins, 1966a), I now

propose that it be replaced by the substitute name,

Echinoderes brevicaudatus.
22 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

11111111111111

100pm

FICURES 26-27.—Echinoderes paciftcus, paratypic female (USNM 5SSS6 RH162.1), neck and

trunk segments: 26, ventral view; 27, dorsal view.
NUMBER 248 23

50um

31

FIGURES 28-31.—Echxnoderes pactficus, segments 12-13: 28, ventral view, lateral half, paratypic

female (USNM 53336, RH162.1); 29, dorsal view, lateral half of same paratypic female; 30,

ventral view, lateral half, paratypic male (USNM 53336, RH162.9); 31, dorsal view, lateral

half of same paratypic male.
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