ROBERT P. HIGGI. Indian Ocean

^ F Kinorhyncha:

1, Condyloderes

and Sphenoderes,

New Cyclorhagid

Genera

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • 1969 NUMBER 14

SMITHSONIAN CONTRIBUTIONS TO

ZOOLOGY

NUMBER 14

Robert P. Higgins India n Ocea n

Kinorhyncha:

1, Condyloderes

and Sphenoderes,

New Cyclorhagid Genera

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ABSTRACT

Higgins, Robert P. Indian Ocean Kinorhyncha: 1, Condyloderes and Sphenoderes,

New Cyclorhagid Genera. Smithsonian Contributions to Zoology, 14:1-13. 1969.—

Condyloderes multispinosus (Mclntyre, 1962), new genus, new combination; Condyloderes paradoxus, new species; and Sphenoderes indicus, new genus, new species

(phylum Kinorhyncha) are described from the coatsts of Scotland and India.

The first kinorhynchs reported from the Indian

Ocean were Campyloderes vanhoeffeni Zelinka, 1913

and Echinoderes ehlersi Zelinka, 1913. The former of

these cyclorhagid species was collected during the

Deutschen Siidpolar Expedition, 1901-1903 at both

Gauss Station, Antarctica (Davis Sea) and Kerguelen Island. A second species, C. macquariae Johnston, 1938, was described from Macquarie Island, and

until C. adherens Nyholm, 1947a was found off the

Swedish Coast, the genus was considered Antarctic in

distribution. Recently C. macquariae has been recorded from the tropical waters of New Caledonia

(Higgins, 1967) but none of the other species have

been found since their original description.

The second Indian Ocean kinorhynch, Echinoderes

ehlersi, is known only from a single collection from

Zanzibar made by Vanhoeffen in 1885 and was the

first kinorhynch to be described from tropical seas

(Zelinka, 1913).

Two additional species of the order of Cyclorhagida,

E. bengalensis Timm, 1958 and E. sonadiae Timm,

1958, were described from the vicinity of Cox Bazar,

East Pakistan. A few years later, an additional cyclorhagid species, Cateria gerlachi Higgins, 1968, was

discovered in a beach at Waltair, India (Ganapati and

Rao, 1962; Rao and Ganapati, 1966) and has been discussed in the first of the current series of papers based

upon the Kinorhyncha I have collected from the

Indian Ocean.

Between 1964 and 1965, German scientists on board

the Meteor found kinorhynchs at record depths of 4690

m off the coast of the Somali Republic (Thiel, 1966)

but no information has been published on the taxonomic identity of the specimens.

During the spring of 1964 I made extensive collections of kinorhynchs along the coasts of India,

Kenya, and the island of Nosy Be, Malagasy Republic.

Robert P. Higgins, Office of Oceanography and Limnology,

Smithsonian Institution, Washington, D.C. 20560.

The report which follows concerns specimens representing new genera of the families Centroderidae Zelinka, 1896, and Semnoderidae Remane, 1936.

METHODS.—Samples of the upper few centimeters

of fine sediment were taken by a meiobenthic dredge.

The sediment was placed in buckets and diluted about

three times its volume with seawater. Minute bubbles

of air were forced through the mud by a hand pump

fitted with a plastic tube and stone air-breaker.

Because of their hydrophobic cuticle, kinorhynchs

brought to the surface by bubbles are retained on the

surface film. By placing a piece of moderately absorbent

paper on the surface film, the trapped meiofauna can

be removed and washed into an accumulating filter

and then preserved by 7 percent formalin.

In the laboratory, specimens were stained with

Semichon's and transferred to a glycerin-alcohol

solution, which was allowed to evaporate to glycerin.

Specimens were removed from the glycerin and individually placed in Hoyer's mounting medium between

two coverslips and positioned in Cobb aluminum slide

frames. This mounting procedure makes it possible to

observe both surfaces of the specimen with oil immersion phase optics.

Each specimen was meristically analyzed and the

resulting data are expressed in a standard format of

abbreviations and terminology (Higgins, 1968).

Measurements are given in microns (/*). Total

length (TL) is measured on the midline, from the anterior margin of segment 3 (the 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 ventral (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 twelfth ventral plates.

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

accessory appendages (LA) are numbered by segment

1
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

and their cumulative mean length expressed by DM,

LM, and LAM. Measurements are given for the lateroterminal spines (LTS), lateroterminal accessory spines

(LTAS), and midterminal spine (MTS).

Various indices using the above measurements are

indicated by the relative abbreviations separated by a

slash (/) and are calculated by dividing the first measurement by the second.

ACKNOWLEDGMENTS.—I would like to thank Dr.

T. S. Satyanarayna Rao, International Indian Ocean

Expedition Liaison Officer, Bombay, for his extensive

help in arranging my research in India; Dr. P. N.

Ganapati and the staff of the Department of Biology,

Andhra University, Waltair, India, and C. R. Eswaran

and T. S. Menon, Fisheries Research Station, Mr.

A. D. Mclntyre, Department of Agriculture and Fisheries for Scotland for his assistance in providing certain

specimens mentioned in this study; Mr. J. W. Cowles,

British Museum (Natural History) for making holotype material available to me; and Drs. H. L. Sanders

and R. R. Hessler, Woods Hole Oceanographic Institution, Woods Hole, Massachusetts, my companions

in this portion of the United States Program in Biology

of the International Indian Ocean Expedition.

This study was made possible by the National Science Foundation's support of the United States Program in Biology, IIOE, and grant GB 3069 which is

gratefully acknowledged. The manuscript was written

while the author was in residence with the SystematicsEcology Program, Marine Biological Labratory, Woods

Hole, Massachusetts.

Key to the Adults of the Kinorhynch Genera

1. First trunk segment entire, bivalvate, or with single midventral plate; round or oval in

cross-section; lateral, dorsal, and terminal spines usually well developed (Order Cyclorhagida) 3

First trunk segment with single, strongly arched dorsal plate, single midventral plate and

two lateroventral plates; trunk spines, if present, restricted to lateroterminal spines (Order

Homalorhagida, Family Pycnophyidae) 2

2(1). With lateroterminal spines Pycnophyes

Without lateroterminal spines Trachydemus

3(1). First trunk segment entire, without midventral plate (Suborder Cyclorhagae) 4

First trunk segment bivalvate, and/or with midventral plate; if not bivalvate, midventral

plates on segments 3-8 7

4(3). First and second trunk segments entire; midterminal spine absent (Family Echinoderidae)

Echinoderes

Second trunk segment with two lateroventral plates; midterminal spine present (Family

Centroderidae) 5

5(4). Elongate spines extending from posterolateral margins of first trunk segment; lateroterminal

accessory spines present in addition to lateroterminal spines 6

Elongate spines not present on posterolateral margins of first trunk segment; lateroterminal

accessory spines absent Condyloderes, new genus

6(5). Lateral spines on most trunk segments; few lateral accessory spines present . . Campytoderes

Lateral spines on few trunk segments, without lateral accessory spines Centroderes

7(3). Midventral plate on segments 3-8, tapering in posterior progression; lateroventral articulation zone weakly developed; middorsal spine absent on segments 3, 7, 9, and 12 (Suborder

Cryptorhagae, Family Cateriidae) Cateria

Midventral plate, if present, restricted to first trunk segment; first trunk segment with bilateral plates; lateroventral articulation zone obvious; middorsal spines on all trunk segments

(Suborder Conchorhagae, Family Semnoderidae) 8

8(7). Midventral and middorsal plate present on first trunk segment; with well-developed placids

(series of plates at anterior margin of first trunk segment) ... . Sphenoderes, new genus

Midventral and middorsal plate absent; without distinct placids Semnoderes

Order CYCLORHAGIDA (Zelinka, 1896) Higgins,

1964

CYCLORHAGAE Zelinka, 18%, p. 197.

CONCHORHAGAE Zelinka, 1907, p. 135 [part].

HETERORHAGAE Lang, 1949, p. 8.

HETERORHAGA Gerlach, 1956, p. 124.

DEFINITION.—Kinorhyncha with variable number

of trichoscalids; second segment (neck) usually with

14-16 well-formed placids or neck plates which form

the closing apparatus; trunk segments round to oval

in cross-section; protonephridia each with single liga-
NUMBER 14

ment; gonopores lateral between segments 12-13;

pharynx musculature forming closed ring; oblique

muscles present in most; armor joints, if present, consisting of dorsal acetabulum and ventral condyle; variously provided with lateral, middorsal, and midterminal spines; surface of trunk segments commonly

covered with minute hairs or denticles, arranged in

distinctive patterns.

Suborder CYCLORHAGAE (Zelinka, 1896)

Zelinka, 1928

NOMOSTOMATA Zelinka, 1907, p. 136 [part].

XENOSTOMATA Zelinka, 1907, p. 136 [part].

CYCLORHAGATA Lang, 1949, p. 8.

CYCLORHAGAEA Chitwood, 1958, p. 943.

DEFINITION.—Cyclorhagida with segment 3 entire,

consisting of a closed ring; segments 4 - or 5-13 each

with a dorsal (tergal) plate lateroventrally articulating with a pair of ventral (sternal) plates.

Family CENTRODERIDAE Zelinka, 1896

MESITODERIDAE Zelinka, 1907, p. 136.

CAMPYLODERIDAE Remane, 1936, p. 347.

DEFINITION.—Cyclorhagae with segments 4-13 divided ventrally to form series of paired ventral plates;

segment 13 tapering terminally to form midterminal

spine; dorsal spines on all trunk segments; lateral spines

and lateral accessory appendages (thought to be adhesive tubes) present on many segments, not restricted

to posterior half of trunk.

Condyloderes, new genus

TYPE-SPECIES.—Centroderes multispinosus Mclntyre, 1962.

DEFINITION.—Gentroderidae with six rows of sealids on segment 1 (head), succeeding rows with 10, 10,

20, 10, 28, and 14—15 scalids; pair of needle-like spinoscalids (row 5) centered anterior to each trichoscalid

(row 6) ; spinoscalids of row 1 with 6-9 setae extending

from base; 16 weakly formed placids with knobby projections; lateral and dorsal spines, variously pilose,

strongly flexible, on segments 3-12; lateral spines of

segment 12 slightly more dorsal than others; lateral

accessory appendages, flattened and wide basally, narrowly tubicolous distally, on segments 4, 7, 10, and 11

(and on segment 6 in one species) ; prominent lateroterminal spines on either side of shorter midterminal

spine; without lateroterminal accessory spines; ventrolateral pores, similar to the base of adhesive tubes

of other genera, on segment 3; prominent conical to

tubicolous intersegmental structures positioned laterodorsally between tergal plates and ventrally on sternal

plates of most trunk segments.

Condyloderes multispinosus (Mclntyre, 1962),

new combination

FIGURES 1-9

Centroderes multispinosus Mclntyre, 1962.

DIAGNOSIS.—Condyloderes with three nearly equal

knobby projections along anterior margin of midventral placid, three larger projections near posterior

margin, one projection near anterior margin and

slightly larger projection near posterior margin of

remaining placids; middorsal and lateral spines pilose;

lateral accessory appendages short, ovoid, on segments

FIGURES 1-2.—Condyloderes multispinosus (Mclntyre),

adult female, topotype, TL 370^, trunk segments: 1, dorsal

view; 2, ventral view.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

4, 7, 10 and 11; intersegmental protuberances of seventh ventral plates missing.

HOLOTYPE.—Adult of undetermined sex, TL ca.

350/*; Loch Nevis (West Coast), Scotland; 101 m;

1962?; British Museum (Natural History) 1962/82.

TOPOTYPES— 1 juvenile, TL 272/*, Higgins K80.1;

1 adult of undertermined sex, TL 336/*, Higgins K80.2;

1 female, TL 352/*, Higgins K81.1; 1 female, TL

370/*, (K81.2); USNM 37459; material provided by

Mclntyre without data.

FIGURES 3-8.—Condyloderes multispinosus (Mclntyre),

adult female, topotype, TL 370^: 3, dorsal view, lateral

half, segments 3-4; 4, ventral view, lateral half, segments

3-4; 5, dorsal view, lateral half, segments 7-8; 6, ventral

view, lateral half, segments 7-8; 7, dorsal view, lateral half,

segments 12-13; 8, ventral view, lateral half, segments 12-13.

REDESCRIPTION.—Topotypic adults, TL 336/*-370/*;

MSW-9 88/*-90/*; SW 65/*-80/*; SW/TL 0.18-0.22.

First segment (Figure 9) with 9 thin, unsegmented

oral styles, ca. 30/* surrounding mouth cone; 6 rows

of scalids, first row with 10 uniformly thick, blunt

scalids, ca. 70/*, generally with 9 setae, 10/*-20/x, near

origin, 2 smaller setae at joint; second row with 10

tapering, pointed scalids, ca. 50/*; third row with 20

tapering, pointed scalids, ca. 40/*; fourth row with 10

slightly more robust, tapering, pointed scalids, ca. 35/*;

fifth row with 28 filiform scalids, ca. 30/*, one pair

centered over each of the 14 trichoscalids of row 6;

trichoscalids, heavily setate, ca. 25/*, centered above

placids (segment 2) except for placids adjacent to

midventral placid.

Second segment (neck) with 16 nearly truncate

placids, poorly cuticularized; midventral placid distinctly larger than others, three knobby projections

along anterior margin (Figure 9), three similar but

slightly larger projections near middle; remaining

placids with single projection in both positions, placids

adjacent to midventral placid slightly narrower than

others.

Third segment (first trunk segment) slightly vaulted

dorsally, especially noticeable along median one-fourth

of segment; posterior border of segment with distinctive pectinate fringe (Figures 3-4), basal half lanceolate in longitudinal section, appearing as a series of

lammelate units, becoming aristate at apex to form

border of setae; short, weakly developed hairs on dorsal

and ventral surface, small, dense patch of very short

hairs midventrally, pattern as illustrated (Figures

1-4); distinctive pores on either side of ventral midline, similar to base of adhesive tubes of male homalorhagids but no tubular extensions apparent; patch

of closely set hairs at base of each lateral and middorsal

spine, spines pilose, lateral spine 28/1,-34/*, middorsal

Spine 24/*-30/*.

Fourth segment with two ventral plates; without

lateroventral pores; spinoid intersegmental protuberances on subdorsal posterior border, a second blunttubicolous protuberance more laterally and a third

protuberance centered at posterior border of each

ventral plate; lateral spine 30/*-34/*; lateral accessory

appendage 15/*-16/*, slightly dorsal to lateral spine,

originating from intersegmental zone of tergal plate

near the lateroventral articulation zone, increasingly

ovoid from point of origin, flat in cross-section,

abruptly tapering, becoming filiform distal one-third
NUMBER 14

TABLE 1.—Measurements (/x) and indices for specimens of Gondyloderes multispinosus

Adult

(Mclntyre, 1962

Fig. 1, p. 506)

Juvenile

K80.1

Adult

K80.2

Adult 9

K81.1

Adult 9

USMN 37459 Adult Range

TL

SW

SW/TL

MSW-9

DM

DM/TL

LM

LM/TL

LAM

LAM/TL

LTS

LTS/TL

MTS

MTS/TL

345

80

0.23

90

49.0

0. 14

45.5

0. 13

14

0.04

220

0.64

80

0.23

272

-

-

-

28.7

0. 10

28.2

0. 10

19

0.07

170

0.63

74

0.27

336

-

-

-

39.9

0. 12

36.7

0. 11

21

0.06

210

0.63

96

0.29

352

65

0. 18

88

44.8

0. 13

40.2

0.11

21

0.06

200

0.57

104

0.30

370

80

0.22

90

44.6

0. 12

41.8

0.11

20

0.05

222

0.60

104

0.35

336-370

65-80

0. 18-0. 23

88-90

39. 9-49.0

0. 12-0. 14

36. 7-45. 5

0. 11-0. 13

14-21

0.04-0.06

200-222

0. 57-0. 64

80-104

0. 23-0. 35

to one-half the length; middorsal spine 28/*-34/t; segment with minute hairs, pattern as illustrated; pectinate fringe as in segment 3.

Segments 5, 6, 8, and 9 similar to fourth but without

lateral accessory appendage; subdorsal intersegmental

protuberance more bluntly tubicolous; L-5 34/i40/i,

L-6 35/X-40/*, L-8 38/*-42/*, L-9 40/*-46>; D-5

28(i-42t<L, D-6 30^34/i, D-8 40/*-52/t, D-9 50^-54^.

Segments 7, 10, and 11 similar to preceding segments but with lateral accessory appendages; LA-7

10ju

FIGURE 9.—Condyloderes

multispinosus (Mclntyre),

adult female, topotype TL

370/t; diagrammatic perspective, lateral half, segments 1

(head) and 2 (neck or

placids) showing midventral

placid and 2 lateral placids

only.

20/1-22/*, LA-10 26(1-28(1, LA-11 16/*-20/*; L-7

36/*-42/i, L-10 42/*-50/*, L-11 50/i-58/*; D-7 38/i-50/i,

D-10 52/X-53/*, D-l l 52/X-60/*; ventral intersegmental

protuberance missing in segment 7.

Segment 12 (Figures 7, 8) without intersegmental

protuberances; lateral spine thin, 40/*-42/t, slightly

pilose, dorsally displaced from lateral margin; middorsal spine 40(i; without lateral accessory appendage.

Segment 13 (terminal segment) without hairs, middorsal, or lateroterminal accessory spines; lateroterminal spine prominent, 210/A-222/I, one-fourth to onethird the. trunk length, slightly pilose at base.

See Table 1 for measurements and indices for specimens of Condyloderes multispinosus.

DISCUSSION.—Several unique features appear in

Condyloderes multispinsosus: (1) the series of flexible

setae at the base of the first row of spinoscalids; (2) a

single pair of needle-like spinoscalids (28 total) centered over each trichoscalid (14 total); (3) knobby

thickenings on the otherwise weakly cuticularized

placids; (4) middorsal and lateral spines with minute

hairs; (5) no lateroterminal accessory spines; and

(6) the presence of the tubicolous intersegmental

protuberances immediately below the pectinate fringe

on the posterior margin of all but the last two trunk

segments.

Condyloderes multispinosus was originally placed in

the genus Centroderes after rejecting the genus Campyloderes because of the lack of especially long lateral

spines on segment 3, and because of the size of the

segment relative to those behind it (Mclntyre, personal communication). Indeed, the choice of assignment of Mclntyre's species to either Centroderes or

Campyloderes would be difficult. Very little is known

about either of these two genera but it is not unlikely

that they may be considered congeneric once careful

studies are made.

Although there as superficial similarities between

Condyloderes multispinosus and Centroderes spinosus
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

(Reinhard, 1881) Zelinka, 1928, these similarities are

minor compared with the unique features used as the

basis for erecting the new genus. Certainly there is

merit in noting that although Centroderes lacks lateral

accessory appendages, C. spinosus does have lateral

spines present only in the positions occupied by the

lateral accessory appendages of Condyloderes multispinosus. Both species have dorsally displaced lateral

spines on segment 12, but the middorsal and lateroterminal accessory spines of the terminal segment of

Centroderes spinosus are absent in Condyloderes

multispinosus.

To complicate matters further, Campyloderes adherens may possess both dorsally displaced lateral spines

on segment 12 and the middorsal spine on the terminal

segment as well as lack lateroterminal accessory spines.

Although this opinion is derived from Nyholm's illustration (Nyholm, 1947a, fig. 3, p. 5) it is contradicted

on page 5 of the text which states: ".. . bears a medium

terminal spine (ts), two short lateral terminal spines

(Its) and two lateral spines (Is)." Furthermore,

Nyholm states (1947a, p. 4) : "Nor are sensory hairs

or adhesive tubes produced." Yet, both Text-figure 1

(p. 3) and Text-figure 3 (p. 5) show "se adhesive tube"

and the illustrated appendage is suggestive of the lateral

accessory appendage in Condyloderes multispinosus.

The illustrated placids of Nyholm's species, however,

are not like those of Condyloderes multispinosus but

are typical of those found in both Centroderes spinosus

and Campyloderes macquariae which I have studied.

In addition to the unique characteristics already

listed for the genus Condyloderes, several contrasting

characteristics of the genera Campyloderes and Centroderes should be mentioned. First of all, the head and

neck in the latter two genera are significantly different

from these areas in the new genus. The mouth styles

for example, are without joints in Condyloderes multispinosus yet are distinctly 2-joined in the two previously

described genera. The lateroventral elongate spines

which originate on the posterior margin of the first

trunk segment and extend the length of several segments of the latter genera are not evidenced in Condyloderes although the two pores on this same segment

suggest some affinity. Finally, the flexible, pilose middorsal and lateral spines of Condyloderes present a

considerable contrast to the straight and rigid, nonpilose spines of Centroderes and Campyloderes.

DISTRIBUTION.—Scotland: Fladen (North Sea),

Lock Nevis (West Coast), and Lock Torridon (West

Coast).

Condyloderes paradoxus, new species

FIGURES 10-20

DIAGNOSIS.—Condyloderes with unequal submarginal knobby projections on midventral placid, central

projection larger than lateral ones, slightly depressed

to accommodate trichoscalid centered above it, three

equal knobby projections near posterior margin; middorsal spines not pilose, lateral spines sparcely pilose at

base only; lateral accessory appendages lanceolate on

segments 4, 6, 7, 10, and 11; tubicolous intersegmental

protuberances present on seventh ventral plate.

HOLOTYPE.—Adult male, TL 246/*.; about 10 km

off shore E of Visakhapatnam (Bay of Bengal), India;

40 m; brown sandy mud; 26 March 1964; col. R. P.

Higgins (RH37.81); USNM 37460.

ALLOTYPE.—Adult female, TL 263/x; as holotype;

(RH37.78); USNM 37461.

PARATYPES.—Four adult males, TL 216/*-264/t; one

adult of undetermined sex, TL 235/*; one juvenile in

FIGURES 10—11.—Condyloderes paradoxus, new species, adult

male, holotype, TL 246/i, trunk segments: 10, dorsal view;

11, ventral view.
NUMBER 14

molt, TL 213/*; as holotype; (RH37 series, retained in

author's collection) ; four adult males, TL 250/*-

282/*; ca. 5 km SE Kakinada Bay (Bay of Bengal),

India; 40 m; soft brown mud; 23 March 1964; col. R.

P. Higgins (RH36 series, retained in author's

collection).

DESCRIPTION.—Holotypic adult, TL 246/*; MSW-9

68M; SW 64/*; SW/TL 0.26.

First segment (Figure 18) with nine, thin unsegmented oral styles, ca. 15/*, surrounding mouth cone;

six rows of scalids, first row with ten uniformly

thick, blunt scalids, ca. 50/*, generally with nine setae,

10/x-20/t, near origin, two smaller setae at base (Figure

19); second row with ten tapering, pointed scalids, ca.

40/*; third row with 20 tapering, pointed scalids, ca.

30/*; fourth row with ten tapering, pointed scalids, ca.

25/i, basal portion elongated along longitudinal axis;

fifth row with 28 filiform scalids, ca. 17/*, one pair

centered over each of the 14 trichoscalids of row six;

trichoscalids, ca. 20/*, not heavily setate, centered above

placids (segment 2) except for placids adjacent to midventral placid, midventral trichoscalid at margin of

midventral placid (Figure 18), other trichoscalids

above placids, extra spinoscalid, ca. 22/i, in normal

trichoscalid position above placid (Figures 18, 20).

Second segment (neck) with 16 nearly truncate

placids, poorly cuticularized; midventral placid distinctly larger than others, with trichoscalid at midmargin, three knobby projections submarginal, central

projection larger, depressed to accommodate trichoscalid, second row of three nearly equal projections

near base; remaining placids with single submarginal

and basal projection, placids adjacent to midventral

placid slightly narrower than others.

Third segment (first trunk segment) slightly vaulted

dorsally, especially noticeable along median one-fourth

of segment; posterior border of segment with distinctive pectinate fringe (Figures 12-13), basal half lanceolate in longitudinal section, appearing as a series of

lammelate units, becoming aristate at apex to form

border of setae; very short, weakly developed hairs on

dorsal and ventral surface, pattern as illustrated (Figures 10—13); distinctive pores on either side of ventral midline, similar to base of adhesive tubes of male

homalorhagids but no tubular extensions apparent;

spinoid intersegmental protuberance on subdorsal posterior border and at posterior border of each ventral

plate; slight indication of closely set hairs at base of

lateral spines, lateral spine 29/t; middorsal spine 28/*.

Fourth segment with two ventral plates; without

lateroventral pores; spinoid intersegmental protuberances on subdorsal posterior border and centered at

posterior border of each ventral plate; lateral spine

28/*; lateral accessory appendage 18/*, slightly dorsal

to lateral spine, originating from intersegmental zone

of tergal plate near the lateroventral articulation zone,

basal two-thirds lanceolate-ovoid, flattened in crosssection, abruptly tapering, becoming filiform distal

one-third; middorsal spine 29/*; segment with minute

5Op

15

16 17

FIGURES 12—17.—Condyloderes paradoxus, new species,

adult male, holotype, TL 246^,: 12, dorsal view, lateral half,

segments 3-4; 13, ventral view, lateral half, segments 3-4;

14, dorsal view, lateral half, segments 7-8; 15, ventral view,

lateral half, segments 7-8; 16, dorsal view, lateral half, segments 12-13; 17, ventral view, lateral half, segments 12-13.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

D

19 20

FIGURES 18-20.—Condyloderes paradoxus, new species, adult male, holotype, TL 246/t: 18,

diagrammatic perspective, lateral half, segments 1 (head) and 2 (neck or placids) showing

midventral placid and 2 lateral placids only; 19, outer surface view, spinoscalid, row 1, showing

basal setae; 20, diagrammatic perspective, anterior view, scalid arrangement, centermost row

1 through row 5 are spinoscalids, outermost row (6) with trichoscalids (note ventral trichoscalid

displacement and extra spinoscalid).

hairs, pattern as illustrated; pectinate fringe as in

segment 3.

Segments 5, 8, and 9 similar to fourth but without

lateral accessory appendages; L-5 31/*, L-8 33/*,

L-9 33/*; D-5 30/*, D-8 31/*, D-9 32/*.

Segments 6, 7, 10, and 11 similar to preceding segments but with lateral accessory appendages; LA6 18/*, LA-7 22/*, LA-10 27/*, LA-11 22/*; L-£ 33/*,

L-7 34/*, L-10 32/*, L-ll 36/*; D-6 29/*, D-7 32/*,

D-10 34**,D-1136*t.

Segment 12 (Figures 7-8) without subdorsal

intersegmental protuberances; lateral spine thin,

dorsally displaced, not pilose, 32/*; middorsal spine

38/*; without lateral accessory appendage.

Segment 13 (terminal segment) without hairs,

middorsal spine, or lateroterminal accessory spines;

lateroterminal spine prominent, 170/*, slightly over

two-thirds the trunk length; midterminal spine 85/*,

nearly one-third the trunk length.

See Table 2 for measurements and indices for

selected specimens of Condyloderes paradoxus, new

species.

DISCUSSION.—The most easily observed feature of

Condyloderes paradoxus, new species, which distinguishes it from C. multispinosus is the former species'

lateral accessory appendage on segment 6. Although

the displaced midventral trichoscalid on the anterior

margin of the midventral placid and the extra spinoscalid in its normal position above the placid are distinctive, the head must be everted in order to observe

these characters.

A close examination of the other placids reveals a

slight modification in this same relative position, thereby suggesting that such a phenomenon may have been

more extensive in the placid row during its development or evolutionary history.

The presence of the midventral trichoscalid on the

border of the enlarged midventral placid corresponds

with the submarginal positioning of the knobby projections typical of the genus. In C. paradoxus, new
NUMBER 14

TABLE 2.—Measurements (/z) and indices for selected specimens of Condyloderes paradoxus, new species

Adult d< Adult 9

Juvenile Adult <? Adult d" USNM Adult tf USNM Adult

RH37.82 RH37.85 RH37.83 37460 RH36.29 37461 Range

Holotypc Allotype

TL

SW

SW/TL

MSW-9

DM

DM/TL

LM

LM/TL

LAM

LAM/TL

LTS

LTS/TL

MTS

MTS/TL

213

-

-

-

28.0

0. 13

23.4

0. 11

16.6

0.08

120

0.57

64

0.30

216

-

-

-

28.4

0. 13

32. 1

0.15

17.6

0.08

162

0.75

96

0.22

245

62

0.25

68

32.3

0. 13

31.7

0. 13

20.0

0.08

155

0.63

86

0.28

246

64

0.26

68

31.9

0. 13

32. 1

0.13

21.4

0.09

170

0.69

85

0.29

282

64

0.23

70

33. 1

0. 12

33.5

0.12

18.6

0.07

160

0.57

broken

263

-

-

-

32.6

0. 12

33.6

0. 13

18.6

0.07

148

0.56

88

0.30

216-282

62-64

0. 23-0. 26

68-70

28.4-33. 1

0. 12-0. 13

31. 7-32. 1

0. 12-0. 15

17.6-21.4

0. 07-0. 09

148-170

0. 56-0. 69

85-96

0. 22-0. 30

species, the central projection is slightly enlarged and

posteriorly displaced. The remaining placid characteristics are similar to those described for C. multispinosus.

In C. paradoxus, new species, there is less hair on

the trunk segments and a corresponding lack of hair

on the spines; the middorsal spines lack hair and the

lateral spines possess only a few minute hairs basally.

There is a vague pattern of hairs displayed on the dorsal and ventral trunk plates which differs from that of

C. multispinosus.

The adults of C. paradoxus, new species, are larger

(TL 216/*-282/x) than those of C. multispinosus (TL

336/i-370/i). The standard width index (SW) of the

former species (0.23-0.26) is greater than in C. multispinosus (0.18-0.23) making the trunk of C. paradoxus, new species, relatively wider than that of C.

multispinosus. Similarly, the average relative length

of the lateral accessory appendages of the new species

(0.07-0.09) is greater than in C. multispinosus (0.04-

0.06), and in addition to being relatively longer, those

of C. paradoxus, new species, are more lanceolate than

in C. multispinosus.

The muscular pharynx was visible in two specimens

of C. paradoxus, new species. This structure is narrowly elongate, 20/* X 80/t. In the cyclorhagid family

Echinoderidae, Echinoderes has a nearly eliptical or

somewhat rounded pharynx. Within the family Centroderidae, Centroderes has a distinctly pear-shaped

pharynx, slightly narrowed anteriorly; in Campyloderes, the anterior portion is narrowly elongate but the

posterior one-third is distinctly broad.

In another suborder, Conchorhagae, Semnoderes

exhibits a nearly oval to pear-shaped pharynx, but in

the suborder Cryptoghagae, the elongate pharynx of

Catena is remarkably similar to that of Condyloderes

paradoxus, new species, in its relative dimensions

(length width ratio of 4:1).

Suborder CONCHORHAGAE Zelinka, 1896

CONCH OR HAG ATA Lang, 1949, p. 8.

DEFINITION.—Cyclorhagida with third segment

bilaterally divided into clamshell-like closing apparatus,

with or without wedgeshaped midventral and middorsal plates; placids weakly developed or with 16

distinct, well-cuticularized placids; mouth styles 2-

jointed; scalids in six to seven rows, trichoscalids present or absent; middorsal spines on all trunk segments,

lateral spines on segments 4 - or 5-12, lateral accessory

appendages on several segments, lateroterminal accessory spines well developed; midterminal spine

usually equal to trunk length.

Family SEMNODERIDAE Remane, 1936

PENTACONTIDAE Zelinka, 1907, p. 135

DEFINITION.—Same as for suborder.
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Sphenoderes, new genus

TYPE-SPECIES.—Sphenoderes indicus, new species.

DEFINITION.—Semnoderidae with 16 well-developed

placids; scalids in six rows, trichoscalid row present;

wedge-like midventral and middorsal plates between

bilateral plates of third segment; middorsal spines on all

trunk segments, lateral spines on segments 5-11, lateral

accessory appendages on segments 7, 10, and 11.

Sphenoderes indicus, new species

FIGURES 21-23

DIAGNOSIS.—Sphenoderes as genus.

HOLOTYPE.—Adult female, TL 256/*; about 10

km off shore E of Visakhapatnam (Bay of Bengal),

India; 40 m; brown sandy mud; 26 March 1964; colR. P. Higgins (RH37.89); USNM 37462.

ALLOTYPE.—Adult male, 256/*; as holotype; (RH

37.91) ; USNM 37463.

PARATYPES.—Two adult females, TL 232/*-240/*;

one adult male, 232/*; as holotype; (RH37 series, retained in author's collection). Two adult females,

TL 226^-280/*; ca. 5 km SE Kakinada Bay (Bay of

Bengal), India; 40 m; soft brown mud; 23 March

1964; col. R. P. Higgins (RH36 series, retained in

author's collection). Two adult females, TL 228/*-

248/*; ca. 12 km off shore from largest tributary of

Coloroon River, near Porto Novo (Bay of Bengal), India; 22 m; brown sandy mud; 14 March 1964; col.

R. P. Higgins (RH27 series, retained in author's collection). One adult male, TL 305/* and one adult

female, TL 400/*; near "Rozi" W Jamnagar (Gulf

of Kutch), India; 6 m; gray-brown mangrove mud;

14 February 1964; col. R. P. Higgins (RH4 series,

retained in author's collection).

DESCRIPTION.—Holotypic female, TL 256/*;

MSW-7 90/*; SW 64/*; SW/TL 0.25.

First segment (head) (Figure 23) with six rows of

scalids; first row with ten uniformly thick, blunt

scalids, ca. 42/* with two setae at joint, 6/*-8/*, second

row with ten tapering scalids, ca. 38/*; third row with

20(?) tapering scalids, ca. 30/*; fourth row with ten

(?) tapering scalids, ca. 23/*; fifth row with ten(?)

tapering scalids; ca. 20/*; sixth row with 14 trichoscalids, weakly ringed, not setose, ca. 13/*; protrusible

proboscis with 7-9 2-jointed oval styles, ca. 20/*.

Second segment (Figures 21-22) with 16 well-developed placids, middorsal and midventral placids with

broad base, adjacent placids narrowly rounded at anterior margin, remaining placids distinctly truncate.

Third segment (first trunk segment) with wedgeshaped middorsal and midventral plates between bilateral plates of third segment; robust middorsal spine

22/*; minute hairs on surface of plates forming slight

pattern, poorly developed anteriorly, becoming more

distinct on posterior margin of segment.

Fourth segment with lateroventral plates articulating

with dorsal plate, lateroventral articulation zone continuous with midventral plate of first trunk segment;

middorsal spine 24/*; pachycycli (thickened anterior

margins) distinct but articulation mechanisms of anterolateral margins of ventral plates not clear; adhesive

tubes not evident; hair pattern similar to third segment;

posterior margins with striate extensions of hair-like

processes.

Fifth segment similar to fourth, middorsal spine 24/*;

with robust lateral spine near lateroventral margin of

dorsal plate, 20/*; dorsoventral muscle scars prominent,

horizontally oriented subdorsally and in middle of

each ventral plate.

FIGURE 21—22.—Sphenoderes indicus, new genus, new species,

adult female, holotype, TL 256^, neck (placids) and trunk

segments, midterminal spine partially shown (MTS 296^) :

21, dorsal view; 22, ventral view.
NUMBER 14

TABLE 3.—Measurements (it) and indices for selected specimens of Sphenoderes indicus, new species

11

Adult 9

RH36.86

Adult 9

USNM

37462

Holotype

Adult 9

RH4.120

Adult cf

RH37.93

Adult c?

USNM

37463

Allotype

Adult o*

RH4.121

Adult

Range

TL

SW

SW/TL

MSW-7

DM

DM/TL

LM

LM/TL

LAM

LAM/TL

LTS

LTS/TL

LTAS

LTAS/TL

MTS

MTS/TL

226

-

-

-

31.5

0. 14

23.5

0.01

15

0.07

76

0.34

38

0. 17

330

1.46

256

64

0.25

90

32.7

0. 13

24.0

0.09

18

0.07

76

0.30

43

0. 17

296

1. 16

400

-

-

-

31.7

0.08

27.5

0.07

18

0.05

80

0.20

42

0.11

376

0.94

232

-

-

-

30.3

0. 13

25.0

0.09

17

0.07

67

0.29

37

0. 16

280

1.21

256

-

-

-

32.5

0. 13

25.6

0. 10

18

0.07

73

0.29

44

0. 17

broken

305

-

-

-

29.0

0. 10

25.6

0.08

18

0.06

broken

44

0. 14

360

1. 18

226-400

-

-

-

29. 0-32. 7

0. 08-0. 13

23. 5-27. 5

0. 07-0. 10

15-18

0. 05-0.07

67-80

0. 20-0. 30

38-44

0. 11-0. 17

280-376

0.94-1.21

Segments 6, 8, and 9 similar to fifth; L-6 21/A, L-8

24/*, L-9 24/t; D-6 28/*, D-8 30/*, D-9 33/*.

Segments 7, 10, and 11 similar to preceding segments

but with lateral accessory appendages; lateral accessory appendages broadly flattened at proximal onehalf, tapering to capillate distally, 18/*; keyhole-shaped

muscle scar(?) immediately anterior to lateral spine

on segment 10 and centered on ventral plates of segment 11; intersegmental protuberance at posterior margin of segment 11; L-7 24/*, L-10 27//,, L-l 1 26/*; D-7

28/*,D-10 35,*,D-1135/*.

Segment 12 with thin middorsal spine, 34/i; lateral

spine, 26/*, ventral muscle scars not evident.

Segment 13 (terminal segment) with thin middorsal spine, 66/*; lateral terminal spine dorsal to lateroterminal accessory spine, 76/*, one-third the trunk

length; lateroterminal accessory spine 43/*, less than

one-fifth the trunk length; gonopores lateroventral

near anterior margin; small posteriorly directed marginal protuberance medial to base of each lateroterminal spine.

See Table 3 for measurements and indices for selected specimens of Sphenoderes indicus, new species.

DISCUSSION.—Because of the strongly curved dorsoventral axis of fixed conchorhagids, dorsoventral orientation of the specimen on a slide mount is difficult.

The problem is further complicated by the relatively

few specimens in the average sample. Consequently,

the chances are great that the specimen will be viewed

laterally. The diagnostic character that must be determined in order to distinguish Sphenoderes indicus, new

species, from Semnoderes is the presence of the middorsal and midventral wedge-shaped plates between the

bilateral plates of the third segment in the former

taxon. This character can be seen by carefully studying

the margins of the anteriormost trunk segment even

when the specimen is mounted laterally.

FIGURE 23.—Sphenoderes indicus, new genus new species,

adult, composite diagrammatic perspective, lateral

half, segment 1 (head).

23
12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

The characteristic placids of Spenoderes may be obscured if the head is retracted, otherwise, no placids

have been seen in the genus Semnoderes. If specimens

are properly cleared, the placids, even when retracted,

are usually visible.

Three species of Semnoderes have been described

and can be distinguished from one another by the number and position of the lateral accessory appendages.

Semnoderes armiger Zelinka, 1928, possesses lateral

accessory appendages on segments 4, 7, 8, 10, and 11;

S. ponticus Bacescu and Bacescu, 1956, has lateral accessory appendages on segments 6, 7, 8, 10, and 11; and

S. pacificus Higgins, 1967, has lateral appendages on

segments 4, 7, 10, and 11.

Semnoderes armiger is well described although the

details of the head are not as elaborately illustrated as

the remainder of the animal; unfortunately, only five

specimens were seen by Zelinka. This species has been

reported from Gullmar Fjord, Sweden by Nyholm

(1947b) and from Fladen and Loch Nevis, Scotland

byMdntyre (1962, 1964).

Semnoderes pacificus is known from New Caledonia

and Redondo Beach, California. This is a particularly

noteworthy species in that the first row of spinoscalids

are distally segmented and the other rows of spinoscalids have medially serrated borders; trichoscalids are

present in this species but not in the others.

The type-material of Sphenoderes indicus, new species, included eleven adult individuals and three juveniles. Of the adults, the two from the Northwest

Coast of India, Bay of Kutch (Jamnagar) are considerably larger (305/x-^00/i) than the average (ca.

250/*) from the East Coast (Porto Novo to Visakhapatnam). The only morphological difference noted was a

more flexible and seemingly segmented middorsal

spine on the twelfth segment of the Jamnagar (RH4

series) specimens. The lateral mounting of the latter

specimens prevents proper doroventral analysis, but

the remaining diagnostic characters seem to be identical with those from the East Coast. I should like to

call the readers attention to Table 3, however, in

order to note the potentially important discrepancies in

the indices for DM/TL, LM/TL, LAM/TL, and

LTAS/TL.

More intensive study of the Jamnagar population

may reveal that two different species exist.

DISTRIBUTION.—Jamnagar (Gulf of Kutch), Porto

Novo, Kakanada, and Visakhapatnam (Bay of Bengal), India.

literature Cited

Bacescu, M., and E. Bacescu

1956. Kinorhinchii—Reprezentanti ai Unei Clase de

Animale, Noua Pentru Fauna Romineasca.

Communications de V Academie de la Republique

Populaire Romaine, 6(4) :543-549, figures 1-6.

Chitwood, B. G.

1958. The Classification of the Phylum Kinorhyncha.

Proceedings of the 15th International Congress of Zoology,

1958:941-943.

Ganapati, P. N., and G. C. Rao

1962. Ecology of the Interstitial Fauna Inhabiting the

Sandy Beaches of Waltair Coast. Journal of the

Marine Biological Association of India, 4(1):44—57,

figures 1-3, tables 1-11.

Gcrlach, S. A.

1956. Uber einen Aberranten Vertreter der Kinorhynchen

aus dem Kustengrundwasser. Kieler Meeresforschungen,

12:120-124, figures 1-15, plates 40-42.

Higgins, R. P.

1964. Redescription of the Kinorhynch Echinoderes remanei

(Blake, 1930) Karling, 1954. Transactions of the

American Microscopical Society, 83(2):243-247, figures

1-3.

1967. The Kinorchyncha of New Caledonia. In Expedition Francaise sur les Recifs Coralliens de la Nouvelle

Caledonia, 2:75-90, figures 1-12.

1968. Taxonomy and Postembryonic Development of the

Cryptorhagae: A New Suborder for the Mesopsammic Kinorhynch Genus Cateria. Transactions

of the American Microscopical Society, 87(l):21-39,

figures 1-25.

Johnston, T. H.

1938. Report on the Echinoderida. Scientific Reports of

the Australasian Antarctic Expedition, 1911-1914, series

C, 10 (7):1-13, figures 1-7.

Lang, K.

1949. Echinoderida. In Odhner, J. H., editor, Further

Zoological Results of the Swedish Antarctic Expedition

1901-1903, 4 (2): 1-22, figures 1-8.

Mclntyre, A. D.

1962. The Class Kinorhyncha (Echinoderida) in British

Waters. Journal of the Marine Biological Association of

the United Kingdom, 44:503-509, figures 1-2.

1964. Meiobenthos of Sublittoral Muds. Journal of the

Marine Biological Association of the United Kingdom,

44: 665-674.

Nyholm, K. G.

1947a. Campyloderes—An "Antarctic" Genus of Echinoderida

off the West Coast of Sweden. Arkiv fur Z00^°£h

39A (13): 1-6, text-figures 1-3, figures l-A, plate 1.

1947b. Studies in the Echinoderida. Arkiv fur Zpologi,

39A (14): 1-36, text-figures 1-22, figures 1-7, 1-9,

plates 1-2.

Rao, G. C , and P. N. Ganapati

1966. Occurrence of an Aberrant Kinorhynch Cateria styx

Gerlach, in Waltair Beach Sands. Current Science, 35

(8):212-213, figures 1-3.
NUMBER 14 13

Reinhard, W.

1881. Ubcr Eckinoderes und Desmoscolex der Umgegen von

Odessa. Zoologischer Anzeiger, 4:588-591.

Remanc, A.

1936. Gastrotricha and Kinorhyncha. In Bronn, editor,

Klassen und Ordnangen des Tierreichs, 4 (sec. 2, pt. 1,

no. 2): vi + 385, figures 1-297.

Thicl, H.

1966. Quantitative Untersuchungen iiber die Meiofauna

des Tiefseebodens. Veroffentlichungen des Institutes

Jiir Meeresjorschungen in Bremerhaven, 2:131-147,

figures 1-3, tables 1-4.

Timm, R. W.

1958. Two New Species of Echinoderella (Phylum Kinorhyncha) from the Bay of Bengal. Journal of the

Bombay Natural History Society, 55(l):107-109,

figures 1-2.

Zelinka, C.

1896. Demonstration von Taflen der Echinoderes—Monographi c Verhandlungen der Deutschen %oologisehen

Gesellschqft 6. Jahresversammlung, Bonn, 197-199.

1907. Zur Kenntnis der Echinoderen. Zoologischer Anzfiger

32(5): 130-136.

1913. Die Echinoderen der Deutschen Sudpolar Expedition

1901-1903. Deutsche Sudpolar Expedition 1901-1903

14 (Zoology, 6):419-436, figures 1-11, plate 39.

1928. Monographic der Echinodera, iv-}-396 pages, 27 plates.

Leipzig: Wilhelm Englemann.

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