Zoologica Scripta, Vol. 12, No. 3, pp. 161-169,1983 Printed in Great Britain

0300-3256183 $3.00+ .OO Pergamon Press Ltd.

The Norwegian Academy of Science and Letters

A New Species of Kinorhynchus (Homalorhagida, Pycnophyidae) from Australia with a Redescription and Range Extension of Other Kinorhyncha from the South Pacific

ROSEMARY BROWN and ROBERT P. HIGGINS

School of Biological Sciences, Macquarie University, North Ryde, N.S. W., Australia, and Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C., U.S.A.

Accepted 14 July 1983

Brown, R. & Higgins, R. P. 1983. A new species of Kinorhynchus (Homalorhagida, Pycnophyidae) from Australia with a redescription and range extension of other Kinorhyncha from the South Pacific.-Zool. Scr. 12: 161-169.

Kinorhynchus phyllotropis sp.n., from Sydney Harbour, is the first species of the phylum Kinorhyncha to be described from Australian coasts. It appears to be most closely related to K . anomalus (Lang, 1953) from the Chilean coast and possibly, but to a lesser extent, to K. spinosus (Lang, 1949) from the Falkland Islands in the South Atlantic. Kinorhynchus anomalus is rede- scribed and, along with the new species, is compared with the South Atlantic species, the two species known from the Northeast Pacific and the remaining four members of this genus from the North Atlantic, Mediterranean Sea and Black Sea. Cateria styx Gerlach, 1956, is reported from beaches north of Valparaiso, Chile, the first report of this mesopsarnmic cryptorhagid kinorhynch from the Pacific Ocean.

Rosemary Brown, School of Biological Sciences, Macquarie Univerisity, North Ryde, N. S. W., 2113, Australia. Robert P . Higgins, Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D. C. , 20560, U.S. A

Introduction

Of the 99 recorded kinorhynch species based on adults and considered identifiable (Higgins 1983), only 16 have been found in the Southern Hemisphere. Kinorhynchs from the Southern Hemisphere were described for the first time by Zelinka (1913) in his report on Kinorhyncha of the German South Polar Expedition of 1901-1903. The cyclorhagid Echinoderes ehlersi Zelinka, 1913, was reported from Zanzibar and a second cyclorhagid, Cam- pyloderes vanhoeffeni Zelinka, 1913, was found in the Davis Sea. Campyloderes macquariae Johnston, 1938, was collected from Macquarie Island by the Australasian Antarctic Expedition of 1911-1914. Since then, this species has been recorded from New Caledonia (Higgins 1967). The Swedish Antarctic Expedition of 1901-1903 collected four species: the cyclorhagid Echinoderes pilosus Lang, 1949, and the homalorhagids Pycnophyes odhneri Lang, 1949, and P . sculptus Lang, 1949, from South Georgia Island and Kinorhynchus spinosus (Lang, 1949) from the Falkland Islands. The Lund University Chile Expedition of 1948-1949 collected kinorhynchs from the Gulf of Ancud, Chile, which included a cyc- lorhagid juvenile, a male Pycnophyes chilensis Lang, 1953, and four specimens of K . anomalus (Lang, 1953). The mesopsammic cryptorhagid kinorhynch Cateria styx Gerlach, 1956, was discovered in the intertidal sand of the high-energy beach at Macae, Brazil, and then also found directly across the South Atlantic in a similar beach at Lobito, Angola (Delamare-Deboutteville 1957), and at

Ubatuba, Brazil (Higgins 1968). More recently C. styx has been encountered at Cachagua Beach and Renaca Beach, both north of Valparaiso, Chile, by Dr. Fernando Alcazar. These latter records are based on specimens sent to one of us (R.P.H.) for identification and are reported here for the first time.

Echinoderes maxwelli (Omer-Cooper, 1957) was col- lected from the brackish water of the Greater Kleinemonde River, South Africa. In 1961 the French Expedition to the coral reefs of New Caledonia collected Campyloderes macquariae, E . newcaledoniensis Higgins, 1967, and a conchorhagid Semnoderes pacificus Higgins, 1967. A neotenic genus of hornalorhagids, Neocen- trophyes intermedius Higgins, 1969, was discovered at Nosy Be, Madagascar and EchinoderespaciJeus Schmidt, 1974, was described from the Galapagos Islands.

The new species described in this paper is the first report of an Australian kinorhynch. It is ascribed to the genus Kinorhynchus Sheremetevskij , 1974 (= Trachydemus Zelinka, 1907), which includes homalorhagid Kinorhyncha without lateral terminal spines.

Methods

Samples of sandy mud were collected from Hunter Bay, Sydney Har- bour, with a Higgins epibenthic dredge. Kinorhynchs were extracted by pumping a stream of fine air bubbles through a suspension of the sediment (Higgins 1964, 1969, 1971). After six hours in Bouin’s fixative, specimens were stored in 70% alcohol. Most specimens were mounted

161 Zoologica Scripta 12

162 Rosemary Brown and R. P. Higgins

la l b

Fig. I . Kinorhynchusphyllotropis sp.n., male.+. Neck and trunk segments. ventral view.-b. Same, dorsal view

in Hoyer's mounting medium for examination of cuticular structures with Zeiss differential interference contrast optics. Several others were used for scanning electron microscope study. These were dehydrated in alcohol and transferred in graduated steps to arnyl acetate. They were then critical point dried, mounted on glass coverslips on a metal stub and coated with a 4 second carbon precoat and a gold-palladium coating. Photographs were made on a Coates and Welter 106B scanning electron microscope.

The data resulting from microscopic analyses are expressed in a standard format of abbreviations and tcrminology (Higgins 1967, 1969). Measurements are given in micrometres (pm). Ratios (e.g. SW/TL) are expressed as a percentage of total length (TL) measured at a the mid-line, from the anterior margin of segment 3 (first trunk segment) to the posterior margin of segment 13 (last trunk segment), exclusive of spines. Maximum sternal width (MSW) is measured at the antero-vent- ral margin of the widest pair of sternal plates as first encountered in measuring each segment from anterior to posterior. Sternal width at segment 12, or standard width (SW), is measured at the anterior margins of the twelfth sternal plates. Segment length (SL) is a measurement of the chord formed by the optical section view of the left (or right) margin of each segment from the anterior edge of the pachycyclus (thickened anterior portion of each segment) to the posterior margin of the segment.

Descriptions of the species

Family Pycnophyidae Zelinka, 1896 Genus Kinorhynchus Sheremetevskij . 1974 (= Trachydemus Zelinka, 1907).

Zoologica Scripta 12

Kinorhynchus phyllotropis sp.n. (Figs. 1-3)

Muterial. Fifty adults from Hunter Bay, Sydney Harbour (Port Jackson), 33"48'30"S., 151"15'24E. Collected from sandy mud (mcan grain size 310 F m ) . 6 m depth, 5 May and 28 Dec. 1980 by R.B. The holotype adult male (W 196 192), allotype adult female (W 196 193) and paratype adult female (W 196 194) are deposited in the Australian Museum, Sydney; paratype adult males and females arc deposited in the National Museum of Natural History. Smithsonian Institution. Washington, DC. G.S.A. (USNM 7400674009). the British Muaeum (Natural History). London (male BM(NH)-1983.1.1: male BM(NH)- 1983.1.2; female 1983.1.3), the Swedish Museum of Natural History, Stockholm (type collection 3216), and the Macleay Museum, Univer\ity of Sydney (male MM-INV 1 : male MM-INV 2: female MM-INV 3 ) .

Diagnoris Anterolateral tergal margins of segment 3 not protruding as horn-like processes; tergal plate of segment 3 with prominent subdorsal setae; mid-dorsal spinose processes extending beyond posterior margins of tergal plates 3-12 with perispinal sensory spots; second segmcnt with four dorsal, slightly indented placids and four even-margined ventral placids; prominent elongate longitudinally oriented cuticular scars on each sternal platc near ventral midline of segments 4-12; unarticulated spinose protuber- ances at lateroterminal margins of tergal plate

Kinorhynchus from Australia 163

2b 2f

2h

2c 2d

Fig. 2. Kinorhynchusphyllotropis sp.n.--a. Segments 2 4 , ventral view, lateral half, male.-b. Same, dorsal view.<. Segments 2 4 , ventral view, lateral half, male.-d. Same, dorsal view.+. Segments 2 4 , ventral view, lateral half, female.-f. Same, dorsal view.-g. Segments 12-13, ventral view, lateral half, female.-h. Same, dorsal view.

Description Adult holotype male (Figs. 1-3) TL715 pm; trunk slightly narrowed anteriorly, more evenly tapered posteriorly, rounded terminally; anterolateral margins of first tergal plate of trunk (segment 3) not protruding as horn-like processes; MSW-8 205 pm, 29 per cent of trunk length; SW 155 pm, 22 per cent of trunk length. Second segment with four dorsal, slightly indented placids and two ventral, even-margined placids (Figs. 2a, b).

Prominent muscle scars situated laterally on tergal plate of segment 3 (Figs. 2b, f), cuticular scars on either side of dorsal midline continuing posteriorly as series on segments 4-12, each located adjacent to pachycyclus, thickened bar of cuticle at anterior edge of segment; crescent-shaped cuticular scars (Figs. 2a, e) centered on episternal plates of segment 3, continuing posteriorly as series, becoming more longitudinally elongate on seg- ments 4-12, each located mesial to midsternal seta.

Trunk segments with prominent, lanceolate rnid-dorsal spinose processes extending beyond posterior margins of tergal plates 3-12 (Figs. lb ; 2b, d , f, h; 3a, b), each process consisting of a ridge of short, dense, cuticular hairs (seen only with scanning electron microscopy) which terminates with a pore (Fig. 3b) before extending beyond tergal margin; process on segment 12 modified, with minute processes extending only slightly on either side of pore (Figs. 2d, h).

Prominent, hollow subdorsal setae (Fig. 3d) approxi- mately 20-25 pm long, on tergal plate of segment 3 anterior to sensory spots (Fig. 3c); second series of shorter

setae about 18-20 pm long, situated laterally near mar- gins of tergal plates 4, 5-7 and 9; third series of setae at lateral margins of tergal plates 4 (male only), 6,8, 10 and 11 and two setae (Figs. 2d, h) at lateral margin of tergal plate 12. [Note: one seta in each pair may belong to the second series but nonetheless gives the appearance of making up a pair near each tergal lateral margin. This reflects the interpretation given for the two setae, near each lateral tergal margin of segment 4 of males (Fig. 2b) but not of females (Fig. 2f)l. Setae centered on sternal plates 5-10; second series near lateral margins of epister- nal plates of segment 3 and sternal plates 5 and 6; males with prominent adhesive tubes (Figs. 2a; 3e), approxi- mately 50-60 pm long, centered on sternal plates of seg- ment 4.

Perispinal sensory spots (Figs. 3a, b) on either side of mid-dorsal spinose processes of tergal plates 3-12; second series of sensory spots subdorsal on tergal plates 3-12 (Figs. 3a, c), those of tergal plate 3 situated posterior to subdorsal setae; third series of sensory spots laterodorsal on tergal plates 3-12, those of tergal plates 3 anteriorly adjacent to prominent lateral muscle scar, those of tergal plate 4 more laterally displaced than preceding or follow- ing sensory spots of same series; fourth series of lateral sensory spots near margins of tergal plates 5-11. Sensory spots (Fig. la) centered on each episternal plate of seg- ment 3, laterally adjacent to crescent-shaped cuticular scars with additional sensory spots situated more post- eriorly, mesial to setae, series continuing with each seta situated centrally, lateral to each cuticular scar of sternal

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164 Rosemary Brown and R. P. Higgins

Fig. 3. Kinorhynchusphyllotropis sp.n., SEM photographs.--a. Posterior section of dorsal midline showing fringed mid-dorsal ridge, slight spinose projection with adjacent sensory spots, subdorsal sensory spots and subdorsal setae of tergal plate on segment 3.-b. Enlarged view of Fig. 3a showing area of spinose projection.+. Subdorsal sensory spot from Fig. 3a.-d. Enlarged view of subdorsal seta from Fig. 3a.-e. Adhesive tube (male) and adjacent sensory spot of left sternal plate of segment 4 . 4 Enlarged view of sensory spot from Fig. 3e.

Zoologica Scripta 12

T 1

100 urn

4a

Kinorhynchus f rom Australia

m 165

w 4b

Fig. 4. Kinorhynchus anomalus, female.-a. Neck and trunk segments, ventral view.-6. Same, dorsal view.

plates 4-12, near lateral margins of sternal plates 11 and 12.

Pachycycli well developed, ventral pachycycli narrow on segment 4, moderate on remaining sternal plates 5-12; dorsal pachycycli prominent, wide, with mesial flanges at dorsal midline (Fig. lb).

Tergal plate of segment 3 with margins devoid of sculpturing, no areas of thin cuticle on anterior margins of episternal or midsternal plates, episternal margin with series of cuticular inclusion spots in some specimens (Figs. l a ; 2a); midsternal plate trapezoidal, lateral margins slightly curved mesially, length 118 pm, basal width 78 pm, 48 pm wide at the apex; segments 4-11 similar in size

and shape; segment 12 roundly tapered; posterolateral border bf terminal tergal plate with prominent, unarticu- lated projections and single small interruption of margin mesial to projections; terminal sternal plates without projections, margins parallel to tergal margin; male with two pairs of penile spines at anterolateral margins of terminal sternal plates; female allotype with single gono- pore in same area.

Males differ from females by presence of adhesive tubes on sternal plates of segment 4, two setae near lateral margins of tergal plate of segment 4 and two pairs of

peni!e spines with simple gonopores near anterolateral margins of terminal sternal plates.

Morphometric data for adult specimens of Kinorhyn- chus phyllotropis are shown in Table 1. The coefficient of variation index shows that measurements least subject to variation within the population sampled are the segment lengths of segments 3 and 9, whereas the most variable measurement is the length of extrusible segment 13.

Etymology. The name of the new species is derived from the Greek words phyllon, ''leaf'' and tropis, "ridge", referring to the foliaceous appearance of cuticular inclusions in the dorsal midline when seen under the light microscope.

Kinorhynchus anomalus (Lang, 1953) (Figs. 4-5)

Trachydemus anomalus Lang, 1953: 6, figs. 5 4 ; Higgins 1961: 86; 1967: 75,1982: 315; 1983: 81. Kinorhynchus anomalus; Sheremetevskij 1974: 985.

Material. Lectotype (here designated-female, permanently mounted for examination) from Station M 29 B, Estero Reloncavi (inner part Bahia Ralun, east of Punta Direccion), 41"24'30"S., 72"19'45"W. Collected from fine sand with mud and terrestrial plant debris at 30-40 m depth on 4 Jan. 1949 (Swedish National Museum of Natural History, type collection 1002 Kinorhyncha 12). Paralectotype (poor condition and not mounted, therefore not examined) from Station M 75B, Canal

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166 Rosemary Brown and R. P. Higgins

Table 1. Meusurements (pm) und indices ( % ) f o r adults of Kinorhynchus phyllotropis sp. n.

Number of Standard Coefficient deviation of variability Character specimens Range Mean

____ TL d d 24 70W330 782 30.4 3.9

99 26 675-760 719 25.0 3.5 d? 50 675-830 749 41.9 5.6

sw d d 24 143-1 63 149 6.8 4.6 ?P 26 143-165 152 6.3 4.1 69 50 143-165 1 50 6.7 4.5

SWiTL d d 24 17-22 19 1.4 7.4 P O 26 20-22 21.1 0.9 4.4 d P SO 17-22 20.1 1.5 7.5

MSW d d 24 205-225 214 6.6 3.1 P P 26 195-235 215 8.4 3.9 d P 5 0 195-235 215 7.6 3.5

MSWiTL d d 24 25-3 1 27 1.3 5.8 P P 26 28-32 30 1.3 4.3 dP 50 25-32 28.7 1.9 6.6

SL-3 di i 24 100-113 109 3.4 3.1 P Q 26 103-1 13 108.5 2.6 2.4 d? 50 100-113 109 3.0 2.8

S L 4 d d 24 65-75 69.8 2.7 3.9 ? ? 26 63-70 66.8 1.9 2.8 d ? SO 63-75 68.2 2.7 4.0

SL-5 5s 24 68-78 74.0 2.9 3.9 P P 26 68-75 72.7 2.2 3.0 d P 50 68-78 73.3 2.6 3.6

SL-6 d d 24 75-85 79.6 2.6 3.3 ?P 26 70-88 77.6 3.6 4.6 d ? 50 70-88 78.5 3.1 4.0

SL-7 Sd 24 75-88 82.5 3.1 3.8 P P 26 75-83 80.3 3.0 3.7 d ? 50 75-88 81 .s 3.2 3.9

SL-8 dS 24 80-93 85.7 3.5 4.1 P Q 26 80-90 83.3 2.5 3.0 d Q 50 8C93 84.5 3.2 3.8

SL-9 d d 24 83-93 87.5 2.7 3.1 P P 26 80-88 84.3 2.0 2.4 d P so 80-93 85.8 2.8 3.3

SL:10 d d 24 83-100 94.1 4.0 5.3 P ? 26 85-95 91.6 2.7 3.0 dP 50 83-100 92.7 3.5 3.8

SL-II d d 24 93-105 98.5 3.4 3.5 9 9 26 88-100 95.0 2.7 2.8 $ 9 5 0 88-1 05 96.6 3.5 3.6

SL-12 $ d 24 93-100 96.5 2.2 2.3 P ? 26 88-100 93.0 3.6 3.9 d ? 50 88-100 94.6 3.5 3.7

SL-12 d d 24 1 8 4 0 33.8 5.2 15.4 P ? 26 15-30 26.8 4.0 14.9 dP 50 1 5 4 0 30.2 5.8 19.2

Moraleda (Penon Blanco), 44"24'00S., 73"34'00''W. Collected from tidal belt rock pools with rather steep rocks o n 24 Feb. 1949 (Swedish National Museum of Natural History, type collection 1002 Kinorhyncha 13).

Diagnosis Closing apparatus of second segment with two slightly indented dorsal mesial placids and two (or four) even- margined ventral placids; anterolateral tergal margins of segment 3 protruding as horn-like processes; tergal plate of segment 3 with prominent subdorsal setae; mid-dorsal spinose processes extending beyond posterior margins of tergal plates 3-12 with perispinal setae on one side (or on both sides) in many anterior segments; triangular to pointed muscle scars anterior to each dorsolateral seta or setal position on tergal plates of segments 4-12; unarticu- lated spinose protuberances at lateroterminal margins of tergal plates of segment 13.

Zoologica Scriptu 12

Redescription of female Lectotype adult female (Figs. 4-5), TL 800pm (about 720 pm according to original description); trunk nar- rowed anteriorly and posteriorly, margins of twelfth seg- ment straight; anterolatcral margins of tergal plate of segment 3 protruding as horn-like processes; MSW-7 214 pm (about 200 pm according to original description), 27 per cent of trunk length; SW 172 pm, 21 per cent of trunk length. Second segment with two slightly indented mesial placids and two (or four?) even-margined ventral placids (Figs. 5e, f).

Prominent muscle scars situated laterally on tergal plate of segment 3 (Fig. 5f), cuticular scars on either side of dorsal midline continuing posteriorly as series on seg- ments 4-12, anterior to setae, not adjacent to pachycycli; crescent-shaped cuticular scars near center of episternal plates of segment 3 (Fig. 5e), continuing posteriorly as

Kinorhynchus from Australia 167

- o \ \

o2 5b

Fig. 5. Kinorhynchus anorna1us.--a. Segments 2-4, ventral view, lateral half, male (after Lang 1953).-b. Same, dorsal view.+. Segments 12-13, ventral view, lateral half, male (after Lang 1953).--d. Same, dorsal view.-e. Segments 2 4 , ventral view, lateral half, female (lectotype).-f. Same, dorsal view.-g. Segments 12-13, ventral view, lateral half, female (lectotype).-h. Same, dorsal view.

series of small circular scars lateral to setae or to setal positions on sternal plates 4-12.

Trunk segments with prominent lanceolate mid-dorsal spinose processes extending beyond posterior margins of tergal plates 3-1 1, reduced on 12 (Figs. 4b; Sb, f).

Prominent subdorsal setae, 2C30 pm long in female (Fig. 5f), possibly longer (up to40 pm?) in male (Fig. Sb), on tergal plate of segment 3, mesial to sensory spots, continuing posteriorly as perispinal setae, present on one or both sides of most anterior mid-dorsal spinose proces- ses of segments 4-11; second series of setae dorsolaterally situated, posterior to cuticular scars on segments 4-11; third series near lateral margins of tergal plates 4-11 with two adjacent setae near margin of tergal plate 12 (Fig. Sh) or one on each tergal plate margin and second setaon each lateral margin of sternal plates (Fig. 5e).

Ventral setae centered on sternal plates 4-9 in female, setae absent on sternal plates 4 of male; second series near lateral margins of episternal plates of segment 3 in female and possibly on lateral margins of sternal plates in male (Fig. Sc); males with relatively short adhesive tubes (Fig. Sa), centered on sternal plates of segment 4.

Perispinal sensory spots (Fig. 4b) on either side of mid-dorsal spinose processes of tergal plates 9 ,11 and 12; second series of sensory spots subdorsal on tergal plates 3-12 beginning with sensory spot posteriorly adjacent to subdorsal cuticular scar on tergal plate of segment 3 , continuing posteriorly, mesial to other subdorsal cuticular

scars and setae; extra sensory spot between perispinal and subdorsal sensory spots of tergal plate of segment 11. Ventral sensory spots near posterior margin of segment 3, posterior to cuticular muscle scar of episternal plates; segment 4 sternal plates of female each with extra sensory spot located mesial to seta; segments 11-12 sternal plates each with prominent sensory spot near ventral midline.

Pachycycli well developed, ventral pachycyli wide on segment 4, moderate on remaining sternal plates but well developed at midline (Fig. 4a); dorsal pachycycli moder- ately wide, without mesial flanges at dorsal midline (Fig.

Tergal plate of segment 3 with margins devoid of sculpturing, no areas of thin cuticle on anterior margins of episternal or midsternal plates; midsternal plate trapezoi- dal, lateral margins straight, length 125 pm, basal width 82 pm, 44 pm wide at apex which flares slightly at margin; segments 4-11 similar in size and shape; segment 12 tapered, lateral margins straight; posterolateral border of terminal tergal plate with prominent unarticulated projec- tions, with single small interruption of margin mesial to projections and fine border fringe; terminal sternal plates without projections, margins roughly parallel to tergal margin; males with two pairs of penile spines (Fig. Sc) at anterolateral margins of terminal sternal plates; female lectotype with single gonopore in same areas.

Males differ from females by presence of adhesive tubes on sternal plates of segment 4, by setae located at

Sf).

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168 Rosemary Brown and R . P. Higgins

adhesive tube positions in female and by possession of two pairs of penile spines near anterolateral margins of termi- nal sternal plates.

Morphometric data for lectotype adult female specimen of K. anomalus: TL 800p.m SW 172pm; SWRL 21%; MSW-7 214pm; MSWiTL 27%; SL-3 118 pm; SL-4 74 pm; SL-5 70 pm; SL-6 82 pm; SL-7 83 p m ; SL-8 85 pm; SL-9 80 pm; SL-10 80 pm; SL-11 92 pm; SL-12 103 pm; SL-13 32 pm.

Discussion

Kinorhynchus phyllotropis sp.n. most closely resembles K. anomalus, but both species share several characters with the only other member of this genus collected from the Southern Hemisphere, K. spinosus.

All three species are of similar dimensions and all have prominent mid-dorsal spinose processes, but each species has characteristics that distinguish it from the other two. In K . spinosus these differentiating characteristics are found on the anterior and posterior aspects of the trunk. The anterior closing apparatus consists of four dorsal placids but only three ventral placids (the only other homalorhagids with three ventral placids are the neotenic Neocentrophyidae). Segment 3, the first trunk segment, has an irregularly denticulate anterior lateral margin and lacks subdorsal setae. The posterior margin of segment 13, the last trunk segment, is pointed and has bulbous latero-terminal spinose projections. In K. phyllotropis distinguishing characteristics include perispinal sensory spots, prominent longitudinally elongate cuticular scars on sternal plates 4-12, a double series of subdorsal sensory spots on tergal plates 4-12 and lateroventral setae near the margim of sternal plates 5 and 6.

Characteristics shared by K. phyllotropis and K. anomalus include a closing apparatus with four dorsal placids and with two ventral placids in K. phyllotropis and two (possibly four) in K. anomalus. Segment 3 of both species has an even anterior tergal margin and subdorsal setae. The location of these setae in relation to the nearby subdorsal sensory spots differs. In K. phyllotropis the setae are situated anterior to the sensory spots, whereas in K. anomalus they are mesial. Both species have subdorsal sensory spots on the segment 3 tergal plate, but K. phyllotropis has a second sensory spot near the anterior margin of the prominent lateral muscle scar. Both species have sensory spots near the posterior margin of segment 3 episternal plates, but K. phyllotropis has a second sensory spot near the prominent lateral muscle scar. Kinorhyn- chus phyllotropis has a double series of subdorsal sensory spots on the tergal plates of segments 4-12. In K . anomalus the lateral series of subdorsal sensory spots is replaced by a series of subdorsal setae on segments 4-12. In bofh K. phyllotropis and K. anomalus the posterior margin of segment 13 is round to truncate and bears unarticulated tapering lateroterminal spinous protru- sions

Kinorhynchus anomalus is distinguished from K. phyllotropis and from K. spinosus by its possession of prominent anterolateral protrusions of the tergal plate of segment 3 and by the straight tapering margin of segment 12, which is evenly curved in K. spinosus. Illustrations of K. spinosus (Lang 1949) show tergal pachycycli with

mesially directed flanges near the midline, which are also seen in K. phyllotropis.

Features of K. phyllotropis which separate it from the remaining six species of Kinorhynchus are the lateral setae formula, the patterns of dorsal and ventral setae and sensory spots and the shapes of the terminal tergal margin and mid-dorsal spinous processes. The remaining species are: K. ilyocryptus (Higgins, 1961) and K. cataphractus (Higgins, 1961) from the northeastern Pacific; K. mainen- sis (Blake, 1930) and K. Zangi (Higgins, 1964) from the northwestern Atlantic; K. giganteus (Zelinka, 1908) from the northern and southern coasts of Europe; and K. paraneopolitanus Sheremetevskij , 1974, from the Black Sea. The description of the latter species is so inadequate that, in our opinion, it must be regarded as species inquirenda.

Other characters distinguish K. anomalus from K. spinosus. Kinorhynchus anomalus has a closing apparatus of two (or four) ventral placids, while K. spinosus has three ventral placids; K. anomalus has an even anterior margin on the tergal plate of segment 3, while in K. spinosus the margin is irregularly denticulate; K. anomalus has prominent anterolateral horns on the seg- ment 3 tergal plate which are reduced in K. spinosus; K. anomalus and K. spinosus have a similar lateral setal formula (differing only in the absence of a seta on segment 4 of K. spinosus); K. anomalus has a truncate terminal tergal margin on segment 13 and this is pointed in K. spinosus.

Kinorhynchus anomalus is clearly distingushed from other members of the genus by the presence of prominent lanceolate mid-dorsal processes and of lateroterminal spinose projections on the terminal tergal margin and by the number and position of setae and sensory spots.

Acknowledgements The authors wish to express their gratitude to Mrs Marie Wallace, research assistant to R.P.H., for her assistance in the preparation of the manuscript, and to Drs Tor Karling and Roy Olerod of the Swedish Museum of Natural History for the loan of the type material of Kinorhynchus anomalus. We are also grateful to the Australian Museum Trust, Sydney, for a post-graduate grant to R.B. which enabled col- laborative field work, to the Sumner Gerard Foundation and to the Smithsonian Fluid Research Fund for their financial support of a portion of this research.

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