Bijdragen tot de Dierkunde, 58 (1): 25-46 — 1988

Taxonomic studies on Dogielinotidae

(Crustacea-) from the Korean coasts

by

Young Won Jo

Institute of Taxonomic Zoology, University of Amsterdam, P.O. Box 4766, 1009 AT Amsterdam,

The Netherlands

Abstract marids of the area often encounter difficulties

with their specimens. As a result, field reports Of forty-eight sandy beach samples from the Korean and leave contained surveys frequently gammarids largely coasts, twenty-two dogielinotid Amphipoda,

four of the Haustorioides unidentified. including new species

1958: H. koreanus H. H. n. latipalpus n. collections of Oldevig, sp., sp., Recently (1982-1986) extensive

n. and H. indivisus n. A key to this nesogenes sp., sp. shallow-water gammaridean amphipods have distribution and of genus, notes on morphology, ecology been made by the author throughout the these four and reconsideration of the species, a genus southern half of the Korean peninsula. The Eohaustorioides Bousfield & Tzvetkova, 1982 are provided. from mud Characters considered of major diagnostic importance samples were taken sandy beaches,

are found in maxilliped, maxilla 1, gnathopods, and flats, gravelly beaches, rocky shores, vary- pereiopods 5, 6 and 7, pleopods, uropods, epimeral plates, ing in depth from medio- to infralittoral (up to oostegites, and telson. 5 48 beaches m deep). Among sandy visited,

found 22 amphipods were at Résumé dogielinotid localities (fig. 1).

Vingt-deux sur quarante-huit échantillons prélevées sur Members of the family Dogielinotidae are des plages de sable des côtes de Corée contenaient des sand-burrowing amphipods endemic to the Amphipodes Dogielinotidés, y compris quatre espèces North Pacific The nouvelles du Haustorioides 1958: H. koreanus (Bousfield, genre Oldevig, region 1970).

H. H. H. is confirmed member of the n. sp., latipalpus n. sp., nesogenes n. sp., et family as a gam- indivisus On donne clé de détermination de n. sp. une ce maridean superfamily Talitroidea (Barnard, des la distribution genre et notes sur morphologie, et 1972; Bousfield, 1982). A total of 7 species in 5 des nouvelles. En on recon- écologie quatre espèces plus, has been described from the North genera sidère le Eohaustorioides Bousfield 1982. genre & Tzvetkova, American Pacific and from the Far Eastern considérés seas Des caractères comme ayant une importance

fournis le la of the diagnostique majeure sont par maxillipède, U.S.S.R. (Bousfield & Tzvetkova, 1982). maxille les les les 1, gnathopodes, péréiopodes 5, 6 et 7, It is remarkable that the diversity of the family pléopodes, les uropodes, les plaques epimérales, les is in so high the western Pacific (10 species oostégites et le telson. including 4 new species), as against only one

species ( Proboscinotus loquax Barnard, 1967) in

INTRODUCTION the eastern Pacific. The proliferation of sand-

burrowing dogielinotids in the western Pacific

world's described marine Of the nearly 4,000 seems analogous to the proliferation of the

19 stricto in the Atlan- gammaridean species (Bousfield, 1981), only Haustoriidae sensu western

species representing 8 families have been tic, with few species along European shores reported from Korean waters (Kim & Kim, (Bousfield, pers. comm.).

Due lack of taxonomic the of 1987). to data, biologists In present study four species and marine Haustorioides 1958 have been ecologists wishing to identify gam- Oldevig, recog-

nized, all of them apparently new to science.

is * The H. koreanus n. most com- Contribution No. 201 of the Institute of Marine large species sp.

beaches the Sciences, National Fisheries University of Pusan, Korea. mon on intertidal sandy throughout

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• 1. Distribution of four of Haustorioides in South Korea = H. koreanus H. H. Fig. species ( n. sp.; * = latipalpus n sp.; ■ =

n. ￿ H. indivisus nesogenes sp.; = n. sp.): 1, Ilkwang; 2, Dadaepo; 3, Nakdong estuary; 4, Chungmu; 5, Samcheonpo;

6, Dolsando; 7, Geomundo; 8, Sinjido; 9, Songhori; 10, Samyang; 11, Hyeopjae; 12, Huksando; 13, Dochodo; 14,

Bigumdo; 15, Byeonsan; 16, Osikdo; 17, Eyeondo; 18, Daecheon; 19, Malipo; 20, Hakampo; 21, Yongyoodo; 22, Baekryeongdo.

southern half of the Korean TAXONOMIC PART peninsula except the east coast, while the other three small

of the in distribu- species genus are allopatric Superfamily TALITROIDEA Bulycheva, 1957 H. tion; latipalpus n. sp. is found only in the Family DOGIELINOTIDAE Gurjanova, 1953

H. indivisus is limited Nakdong estuary, n. sp. Genus Haustorioides Oldevig, 1958 to the west and H. is coast, nesogenes n. sp. Oldevig, 1958: 343; Gurjanova, 1962: 429 (in part); distributed in the region between the two Barnard, 1967: 287; 1969: 209; 1972: 178; Bousfield & species (southwestern islands of Korea). Tzvetkova, 1982: 87. The material is preserved in the Zoologisch Species content. — H. munsterhjelmi Oldevig, 1958 Museum, Amsterdam and the Institute (ZMA) H. japonicus Kamihira, 1977 of Marine Sciences, Pusan H. Bousfield, 1982 (IMS). magnus

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H. gurjanovae Bousfield, 1982 uncleft, tapering distally; maxilliped, outer lobe not

H. koreanus of normal n. sp. reaching outer tip palp segment 1, palp

H. H. indivisus latipalpus n. sp. n. sp.

H. — basis with anterodistal nesogenes n. sp. Gnathopod 1, 2-3 setae only;

H. indivisus 5 n. sp. pereiopod 5, dactylus short; pereiopod 7, segment

with 3 tall longer than segment6; uropod 1, peduncle

— Rostrum medium distinct, telson cleft about 1/4 Diagnosis. spines; at apex; maxilliped, outer

small. medium medium ovoid. lobe of Eye or small, long, extending to outer tip palp segment 1,

short and wide H. Antenna 1 shorter than 2, peduncle of both palp latipalpus n. sp.

3. Pereiopods 6 and 7, segment 6 with 2-3 clusters of antennae short. Mandibular molar strongly setae posteriorly; pereiopod 7, posterior margin of triturating, incisor well toothed, left lacinia 4-5 segments 4 and 5 weakly setose; uropod 1, peduncle dentate. Labium with small inner lobes. Inner with 5-6 tall spines 4

lobe of maxilla 1 short and 2-3 — 6 with 5-7 clusters of small, bearing Pereiopods 6 and 7, segment

setae pereiopod 7, posterior margin of plumose setae on apex, outer lobe with 9 long posteriorly; and 5 4 strongly setose; uropod 1, peduncle pectinate spines, palp minuteand 1-segmented. segments with 9-15 tall spines 6 Dactylus of maxillipedal palp with modified or 4. Gnathopods weakly sexually dimorphic (in male little lacking unguis. longer than in female); pereiopods 6 and 7, dactylus Coxal plates moderately large, first not weakly setose (3-5 setae); epimeral plate 2,

much shorter than reduced, fourth excavate posteriorly. Male posterodistal corner slightly acute, that ofepimeral plate 3; telson lobes apically separated gnathopod 2 larger than 1 and sexually dimor- by distinct notch H. gurjanovae Bousfield, 1982 phic, female gnathopod 2 subequal to — Gnathopods strongly sexually dimorphic; pereiopods 6 gnathopod 1 and different in structure. scarcely and 7, dactylus strongly setose (more than 8 setae); 5-7 in Pereiopods increasing length con- epimeral plate 2, posterodistal corner strongly or

and moderately produced, as in epimeral plate 3; telson secutively, medium or strongly spinose hardly cleft 5 setose, segments 2, 4, 5 of P5-P6 strongly 5. Pereiopods 6 and 7, dactylus with 7-16 setae expanded posteriorly. 4 shorter anteriorly; pereiopod 7, segment distinctly Posterodistal corner of epimeral plate 2 than segment 5; uropod 2, peduncle with 2 slender slightly or strongly produced, that of plate 3 spines and 2 stout spines; pleopods, peduncle with 5-6

telson cleft retinacula; slightly H. n. strongly produced. Rami of uropods 1 and 2 .. nesogenes sp.

— Pereiopods 6 and 7, dactylus with 20-50 setae ante- unarmed. Uropod 3 minute, with 1-3 apical riorly; pereiopod 7, segments 4 and 5 subequal; setae, lacking rami. Urosomites free, 1 dorsally uropod 2, peduncle with 5-6 slender spines only;

2 and 3. Telson entire or overhanging slightly pleopods, peduncles with 10-13 retinacula; telson lobes cleft medially. fused entirely H. japonicus Kamihira, 1977

6. Coxal plates 1-4, lower margins with long setae;

1/2 pereiopod6, segment 5 sublinear (width = length); KEY TO THE WORLD SPECIES OF THE GENUS uropod 2, peduncle with 1 stout spine and 5-6 slender HAUSTORIOIDES

spines; epimeral plate 3, posterodistal corner

1. of with H. koreanus Uropod 2, margin peduncle stout spines only; moderately produced n. sp.

than — Coxal lower with pereiopod 6, segment 5 sublinear, slightly wider plates 1-4, margins mainly short

claw of with thick 5 segment 6; maxilliped, palp spine, setae; pereiopod 6, segment broadly expanded prox- about 2/3 of inner margin of palp segment 2 convex; maxilla 1, imally (width length or more); uropod 2,

palp lacking apical seta 2 peduncle with 8-12 slender spines only; epimeral plate

— Uropod 2, margin of peduncle with slender spines and 3, posterodistal corner strongly produced, length of

5 less than basal width 7 stout spine(s); pereiopod 6, segment more or process greater

broadened; maxilliped, claw of palp lacking unguis, 7. Pereiopod 6, segment 5 broadly expanded proximally,

with slender inner of width about 2/3 of with 14- numerous setae, margin palp seg- length; uropod 1, peduncle

ment 2 vertically straight; maxilla 1, palp with an 15 slender spines; gnathopod2, posterodistal angle of

apical seta 3 palm with 7 spine-teeth; pleopods, peduncle with 14-

2. Gnathopod 1, basis with anteroproximal and 17 retinacula; oostegites on gnathopod 2 subquadrate

anterodistal H. 1982 setae; pereiopod 5, dactylus normal; magnus Bousfield,

5 shorter than — 5 sub- pereiopod 7, segment segment 6; Pereiopod 6, segment very broadly expanded,

uropod 1, peduncle with 3-4 short spines; telson quadrate, length equals width; uropod 1, peduncle

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with 9-12 slender gnathopod 2, posterodistal 1 of 1 spines; Peduncle segment antenna (fig. 2c) of with few weak angle palm spines; pleopods, pedun- twice twice wide as long and as as segment 2; sub- cle with 10 retinacula; oostegites on gnathopod 2 and 3 with bundles of and segments 2 long thick triangular H. munsterhjelmi Oldevig, 1958 (sub)distal setae. Flagellum 10- to 11-

each segment posterodistally armed Haustorioides koreanus segmented; n. sp. with bundle of each setae; 1 or 2 aesthetascs on (Figs. 2-4) first and last segment except one.

Material examined. — Pusan, Dadaepo; 5 Apr. Antenna 2 (fig. 2d) with rather slender 1985, 1 C holotype (ZMA Amph. 108.331a), 1 ovigerous peduncle. Flagellum of 9-10 segments; each 9 allotype (ZMA Amph. 108.331b), 43 ovigerous 99 armed with setae. and 44 20 segment distally long CO" paratypes (ZMA Amph. 108.331c), Labrum with 2 ofmid-medial ovigerous 99 and 22 0"o* paratypes (IMS); 17 Dec. 1982, (fig. 2e) groups

67 4 Mar. 118 17 specimens; 1983, specimens; Apr. 1983, setae; free margin rounded, with setules. 79 11 Oct. 22 Dec. specimens; 1984, many specimens; Mandible with and (fig. 2f): Right 6 strong 1984, 170 specimens; 7 Mar. 1985, 42 specimens. fine teeth lacinia followed many on mobilis, Pusan, Nakdong estuary, Okryudeung; 16 Nov. 1983, proximad by 4 plumose setae; pars molaris with 4 specimens; 20 Jan. 1984, 2 O"O"; 12 Apr. 1984, 2 CO" with and 3 ovigerous 9 9 • two long setae. Left lacinia robust, armed

Prov. 23 Mar. 1 and 1 Kyeongnam, Ilkwang; 1986, O" 5 teeth, followed proximad by 6 plumose setae; ovigerous 9- molaris with 2 pars setae. Prov. Kyeongnam, Chungmu; Sept. 1986, 5 Labium (fig. 2g) with small, but distinct specimens. inner lobes; anterior side of left outer lobe with Prov. Kyeongnam, Samcheonpo, Namildae, 26 May

1986, 5 crcr and about 600 juvs. groove on outer margin.

Prov. Cheonnam, Dolsando, Bangjukpo; 6 Aug. 1986, Palp of maxilla 1 (fig. 2h) 1-segmented, with about 320 juvs. 1 which is than itself. apical seta, longer palp Prov. Cheonnam, Geomundo, Seodori; 23 July 1986, Outer lobe with 9 denticulate spines. Innerlobe 46 juvs. slender, armed with 2-3 plumose setae. Prov. Cheonnam, Sinjido, Myeongsasimri; 11 May

1986, 8 CfC, 16 99 (13 ovigerous) and 138 juvs. Maxilla 2 (fig. 2i) of the usual structure.

Prov. 8 19 Cheonnam, Haenam, Songhori; Aug. 1986, Inner and outer lobes armed with distal and juvs. mediodistal setae; inner margin of inner lobe Prov. Cheonnam, Huksando, Jinri; 22 July 1986, 106 with 4 strong plumose setae. juvs. Inner lobe of with Prov. Cheonnam, Bigumdo, Gurimri; 19 Oct. 1986, maxilliped (fig. 2j) distally three robust dentiform 181 specimens. spines, placed at regular

Prov. 7 10 Cheju, Samyang; Aug. 1986, juvs. intervals; inner margin with 5 plumose setae. Prov. Cheju, Hyeopjae; 8 Aug. 1986, 153 juvs. Outer lobe short, not reaching outer tip of palp Prov. Cheonbuk, Byeonsan; 19 Oct. 1986, 3 99- 1. 2 with segment Palp segment an additional Prov. Cheonbuk, Okgu, Osikdo; 24 July 1986, 162

row of setae inner juvs. submarginal vertically, Prov. Chungnam, Boryeong, Eyeondo; 4 Feb. 1983, margin vertically straight; dactylus linear, with

slender many specimens. apical setae, lacking unguis.

Prov. 4 Feb. 2 and Chungnam, Daecheon; 1983, ccr 1-4 Coxal plates (figs. 3a-d) with numerous 3 99; 9 May 1986, 26 CO" and 20 99 (18 ovigerous). longish setae on lower margin. Prov. Chungnam, Malipo; 21 Oct. 1986, 33 crcr and Gnathopod 1 (fig. with 3 99- 3a), carpus posterior base of lobe Prov. Kyeongki, Baekryeongdo, Gobongpo; 18 Nov lobe; lined with heavy setae.

1986, 3 CTCT and 4 99- Palmar angle of propodus with 4-5 spine-teeth.

Dactylus short, stout. Description. — Body length (without anten- Gnathopod 2 (fig. 3b) strongly subchelate; and male female nae telson): 11.0-12.0 mm, length of propodus about 1.8 times that of 12.5-14.0 Male in mm. habitus as fig. 2a. gnathopod 1. Carpus with long posterior lobe;

Male: Lateral base of cephalic lobes (fig. 2b) acute. of lobe with row dense, short setae. Eyes of medium size, ovoid.

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2. Haustorioides koreanus 11.5 mm from lateral of b, head Fig. n. sp. (￿ holotype Dadaepo): a, aspect habitus; (scale AC);

antenna 2 labrum f, left and mandible (AD); labium (AD); h, maxilla c, antenna 1 (AB); d, (AB); e, (AD); right g,

1 (AD); i, maxilla 2 (AD); j, maxilliped (AC).

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3 4 and Pereiopods and (figs. 3c, d), merus Oostegites on gnathopod 2 (fig. 4b) tapering carpus heavily setose posteriorly. Propodus distally, subtriangular; those on P3-P5 (figs. 4c-

armed with 3-4 pairs of thick setae on e) subquadrate, distally widened. posterodistal half.

Remarks. — This species is closely related to Pereiopod 5 (fig. 3e) densely setose; merus the large species H. munsterhjelmi Oldevig, 1958, slightly longer than wide; propodus with from the Island of Sachalin, U.S.S.R., but it dac- numerous setae along posterior margin; differs from it in the carpus of P6 (sublinear in tylus with 0-1 seta on anterior margin. H. koreanus, in H. 6 and subquadrate munsterhjelmi), Pereiopod (fig. 3f), merus carpus the number the of of spines on peduncle uropod densely setose, not heavily expanded (carpus 1 (14-17 in H. koreanus, 9-12 in H. about twice shorter munsterhjelmi), as long as wide); carpus the less strongly produced of than armed with posterodistal point propodus. Propodus 6 groups epimeral plate 3, and the absence of antero- of with 8-11 setae on posterior margin; dactylus distal setae the basis of 1 and 2 anterior on gnathopods setae on margin. in both 7 sexes. Pereiopod (fig. 3g) moderately setose; From H. Bousfield, 1982, the shorter than magnus present carpus propodus; posterior margin species differs in the less strongly produced of propodus armed with 5-6 bundles of setae; posterodistal point of epimeral plate 3, the dactylus with 21-22 anteromarginal setae. sublinear of the number of carpus P6, Pleopodal peduncles (fig. 3h) each with 9-10 retinacula on the peduncles of the pleopods (9- retinacula; exopodite of first pleopod 26- to 28- 10 in H. koreanus, 14-17 in H. magnus) and the segmented. shape of the oostegite on gnathopod 2 (sub- Epimeral plates 1 and 2 (fig. 3i) with fine in H. koreanus, in H. setules anteroventral Plate triangular subquadrate on their part. 1 with

rounded 2 magnus). slightly posterodistal corner; plate Morphological differences among popula- with small point; plate 3 with medium-sized tions of this small. species are very However, point. show less setation the juveniles strong on Uropod 1 (fig. 3j) with 14-17 slender pedun- appendages, feeble subchelation of gnathopod 2 cular spines (9-10 on outer margin). in the male, a wide merus of P5, and a more Uropod 2 (fig. 3k) with 5-7 longish pedun- arrangement of the three robust teeth- cular but the distal irregular spines, most one on outer in spines on the inner lobe of the maxilliped, as margin short and stout. the three small of Korean Haustorioides. of species Apex uropod 3 (fig. 3-1) with a single seta; But juveniles of this species can be distin- length of apical seta much shorter than half of guished from other Korean species of peduncle. Haustorioides by the shape of the telson, the Telson (fig. 4f) wider than long, cleft for of dense of setules the base of presence a row at about 1/4 of its length; each lobe with 4 the carpal lobe of gnathopods 1 and 2, the slight laterodistal setae, three of which feathered. acumination of the posterodistal corner of

epimeral plate 2, and the additionalrow of sub- Female: Lower margin of coxal plates 1-4 of the marginal setae on palp segment 2 max- than in male. (figs. 4a, b) more strongly setose illiped. Gnathopod 1 (fig. 4a), propodus relatively

with small, 2 slender spine-teeth on palmar Etymology. — The specific name koreanus angle; dactylus slender, long, with 2 blades on refers to the peninsula of Korea where this the tip. species is widely and abundantly distributed. Gnathopod 2 (fig. 4b) subequal to the first.

Lobe of — carpus more elongate; propodus much Distribution and ecology. This species smaller than in male. is one of the most common amphipods on inter-

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3. Hauslorioides koreanus from Fig. n. sp. (￿ holotype 11.5 mm Dadaepo): a, gnathopod 1 (scale AB); b, gnathopod 2

3 4 5 6 7 (AB); c, pereiopod (AB); d, pereiopod (AB); e, pereiopod (AB); f, pereiopod (AB); g, pereiopod (AB); h, pleopod 1 (AB); i, epimeral plates 1 to 3 (AB); j, uropod 1 (AC); k, uropod 2 (AC); 1, uropod 3 (AD).

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4. Haustorioides koreanus 13.0 11.5 from Fig. n. sp. (a-e, ￿ allotype mm; f, ￿ holotype mm, Dadaepo): a, gnathopod

2 with 1 ￿ (scale AB); b, gnathopod oostegite ￿ (AB); c, oostegite of pereiopod 3 ￿ (AB); d, oostegite of pereiopod

4 ￿ (AB); e, oostegite of pereiopod 5 ￿ (AB); f, telson ￿ (AC).

tidal sandy beaches throughout the southern medium-coarse with shell debris). halfof the Korean the At the the peninsula (fig. 1) except type-locality, Dadaepo, highest

of where action is 2 reached around the east coast (Sea Japan), wave density (359/m ) was

midlittoral and the fluctuation of strong and the tidal range is very small (20-30 zone monthly

this is From cm at spring tide). In Chejudo species salinity was 2.73-33.34%o (Jo, 1983). found in the sheltered northern of the this have only part year-round sampling species seems to island, indicating that the prefers pro- a life span of one year, ovigerous females being

found in tected areas. In most sampling localities the spring (March-June), with a large

number of The 13.0 substratum consists of fine to medium sands eggs. allotype (body length

in Daecheon and the sand is carried 181 size 0.37 0.44 (but Baekryeongdo mm) eggs, egg x mm.

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Haustorioides and second from the inside close latipalpus n. sp. one together,

the lateral from the inner (Figs. 5-7) one apart others);

margin with 3 plumose setae. Outer lobe

Material examined. — Pusan, Nakdong estuary, distally tapering, extending to level of outer tip Baekhabdeung; 3 Apr. 1984, 1 cr holotype (ZMA Amph. 1. inner of palp segment Palp short, wide; 108.332a), 1 ovigerous 9 allotype (ZMA Amph. of 2 thick margin segment convex; unguis very 108.332b), and 1 ovigerous 9 paratype (ZMA Amph. and 108.332c); 19 Jan. 1984, 5 crcr , 6 99, and 3 juvs. long.

7 3 Coxal 1-4 paratypes (ZMA Amph. 108.333a); Jan. 1985, CTO" plates (figs. 6a-d) sparsely setose

and 1 9 paratypes lower of (IMS). on margin; anterodorsal corner plate 1 12 Pusan, Nakdong estuary, Okryudeung; Apr. 1984, convex. 1 9; 25 Oct. 1984, 2 99 (1 ovigerous); 7 Jan. 1985, 6 crcr Gnathopod 1 (fig. 6a), anterior margin of and 11 99- basis with 2 distal Base of lobe 1 1 setae. carpal Pusan, Nakdong estuary, Myeongji; May 1984,

ovigerous 9; 8 Jan. 1985, 1 9- with two setules only. Palmar angle of propodus

1984, Pusan, Nakdong estuary, Eulsukdo; 13 Aug. with 2 slender, bifid spine-teeth. 3 99 (1 ovigerous); 23 Oct. 1984, 1 C. Gnathopod 2 (fig. 6b), anterior margin of Pusan, Nakdongestuary, Jinwoodo; 21 Jan. 1985, 1 O" basis with 3 distal setae. Base of carpal lobe and 1 9- with setules in 1. of two as gnathopod Length

propodus about 1.4 times that of gnathopod 1.

Description. — Small species; largest male Pereiopods 3 and 4 (figs. 6c, d), merus and

female 7.8 setose 6.6 mm long, largest mm. carpus moderately posterodistally.

Pereiopod 5 (fig. 6e) moderately setose;

wider than of Male; Lateral cephalic lobes (fig. 5a) rounded merus long; posterior margin pro-

with short. triangular, not exceeding tip of rostrum. Eyes podus a single seta; dactylus

ovoid. 6 about of small, Pereiopod (fig. 6f), carpus linear,

Antenna 1 2 and of (fig. 5b), peduncle segments equal length as propodus; posterior margin

3 with with bundles of long setae on their distal propodus a single seta; dactylus with 1-2

7- each anterior margin. Flagellum to 8-segmented; seg- setae on margin.

short and with few 1 7 ment wide, short setae; or Pereiopod (fig. 6g) poorly setose; merus

aesthetascs each first and shorter than 2 on segment except distinctly carpus; carpus slightly

last than of one. longer propodus; posterior margin pro-

Antenna 2 with slender with of with 4- (fig. 5c) gland cone. podus two groups setae; dactylus

Flagellum 8- to 9-segmented, poorly setose. 6 anteromarginal setae.

Labrum (fig. 5d) without mid-medial setae; Pleopodal peduncles (fig. 5j) each with 5-6

free margin slightly emarginate. retinacula; exopodite of first pleopod 18-

Mandibles (fig. 5e) asymmetrical; right man- segmented.

dible with finely toothed lacinia mobilis; left Epimeral plate 1 (fig. 6h) with small

lacinia mobilis with 4-5 robust teeth. posterodistal point; plates 2 and 3 each with

Labium (fig. 5f) with distinct inner lobes; left medium-sized point.

Peduncle of outer lobe without groove. uropod 1 (fig. 5k) with 3 slender

Palp of maxilla 1 (fig. 5g) vestigial, 1- spines.

segmented, without apical seta; inner lobe Uropod 2 (fig. 5-1) with 4 stout peduncular armed with 2-3 setae only. plumose on apex. spines

of Maxilla 2 (fig. 5h) of the usual structure; Apex uropod 3 (fig. 5m) with 2-3 setae;

inner of inner lobe with of half that of margin 1 strong length setae about peduncle.

plumose seta. Telson (fig. 5n) cleft for about 1/4 of its

with 3 dentiform each lobe 4 three Maxilliped (fig. 5i) strong length; with laterodistal setae, spines of inner lobe arranged irregularly (first of which feathered.

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5. Haustorioides 6.6 from head Fig. latipalpus n. sp. (￿ holotype mm Baekhabdeung, Nakdong estuary): a, (scale AB);

1 2 labrum mandibles labium maxilla 1 h, maxilla b, antenna (AB); c, antenna (AB); d, (AC); e, (AC); f, (AC); g, (AC);

2 (AC); i, maxilliped (AC); j, pleopod 1 (AB); k, uropod 1 (AB); 1, uropod 2 (AB); m, uropod 3 (AC); n, telson (AC).

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6. Haustorioides 6.6 from Fig. latipalpus n. sp. (￿ holotype mm Baekhabdeung, Nakdong estuary): a, gnathopod 1; b,

1 gnathopod 2; c, pereiopod 3; d, pereiopod 4; e, pereiopod 5; f, pereiopod 6; g, pereiopod 7; h, epimeral plates to

3. Scales: all AB.

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7. Haustorioides 7.8 mm from Fig. latipalpus n. sp. (￿ allotype Baekhabdeung, Nakdong estuary): a, gnathopod 1; b,

with of of of 5. Scales: gnathopod 2 oostegite; c, oostegite pereiopod3; d, oostegite pereiopod 4; e, oostegite pereiopod

all AB.

Female: Gnathopod 1 (fig. 7a) similar to that the posterodistal point of epimeral plates 2 and of and and 3. male, but propodus dactylus thinner; 3, the longish setae on uropod

with minute In the the dactylus long, unguis a blade. particular, present species shares

Gnathopod 2 (fig. 7b) similar to gnathopod following characters with H. indivisus: peduncle

1; propodus triangular; dactylus slender. of uropod 2 with stout spines only, unguis of

and of maxilla 1 bare. Oostegites on P3-P5 (figs. 7c-e) subquadrate, maxillipedal palp palp

that of distally widened; gnathopod 2 (fig. 7b) However, the present species is well- tapering, pointed. characterized by its cleft telson (about 1/4),

of absence of anteroproximal setae on the basis

Remarks. — This species is similar to H. gnathopod 1, and the peculiar structure of the

and H. indivisus in the short and nesogenes relatively poor maxilliped (elongated outer lobe, setation of the appendages, about equal size of wide palp).

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— lacinia mobilis Etymology. The proposed specific name, Mandible (fig. 8e): Right latipalpus (from the Latin, meaning "wide bifid, with fine teeth, followed proximad by 4 armed with palp"), alludes to the palp of the maxilliped plumose blades. Left lacinia mobilis

followed blades. (especially segment 2) which is rather short and 4 robust teeth, by 6 plumose wide. Labium (fig. 8f) with inner lobes; left outer

lobe grooved. Distribution and ecology. —The species, Maxilla 1 (fig. 8g), palp 1-segmented, with 1 col- probably a brackish-water endemic, was inner apical seta; lobe with 2-3 plumose setae. in the Five lected Nakdong estuary (fig. 1). Maxilla inner and 2 (fig. 8h), setation on sand dunes where the species occurs are all each outer lobes extended to lower halfof lobe; In situated within a radius of 3 km. Okryu- of inner lobe inner margin with 3 strong with deung, the purest sandy bottom, this plumose setae. H. koreanus. species was accompanied by Maxilliped (fig. 8i) with 3 robust dentiform The animal inhabits intertidal sand or spines of inner lobe arranged irregularly; inner muddy sand. The salinity of the interstitial margin with 4 plumose setae. Outer lobe not water fluctuates extremely during the tides and distal end of reaching outer palp segment 1. by outflowing river water. During the study Inner margin of palp segment 2 straight; dac- period ovigerous females were found in April, with bundle of slender tylus linear, a setae. October with small May, August, and 1984, a Coxal plates 1-4 (figs. 9a-d) poorly setose on number of The 7.8 eggs. allotype (body length lower margin; proximal part of plate 4 excavate mm) carried 18 eggs, egg size 0.29 x 0.37 mm. posteriorly and sloped anteriorly.

Gnathopod 1 (fig. 9a), anterior margin of

basis with 3 long setae and 6 small distal setae.

lobe with basal setules Palmar Haustorioides Carpal 2-3 only. nesogenes n. sp. of swollen, armed with 2 (Figs. 8-10) angle propodus

slender, bifid spine-teeth; claw short, stout,

Material examined. — Prov. Cheonnam, Sinjido, without unguis. Myeongsasimri; 11 May 1986, 1 cr holotype (ZMA Gnathopod 2 (fig. 9b), anterior margin of Amph. 108.334a), 1 ovigerous 9 allotype (ZMA Amph.

and 4 era* basis with 3 distal lobe with 2 basal 108.334b), 4 99 (3 ovigerous) paratypes (ZMA setae; carpal

4 and 4 Amph. 108.334c), 99 (3 ovigerous) crty para- setules; length of propodus 1.5 times that of (IMS). types gnathopod 1. Prov. Cheonnam, Dochodo, Simok; 6 Oct. 1986, 4 crcf Pereiopods 3 and 4 (figs. 9c, d), merus and and 3 99-

setose Prov. Cheonnam, Bigumdo, Gurimri; 19 Oct. 1986, carpus moderately posterodistally. 5 322 specimens. Pereiopod (fig. 9e) moderately setose;

much wider than merus long; posterior margin

— male 6.7 Description. Body length: mm, with of propodus with a single seta; dactylus 0-2 female 9.2 mm. anterior setae.

Male: Lateral cephalic lobes (fig. 8a) rounded Pereiopod 6 (fig. 9f), merus expanded; triangular, extending to about the tip of the carpus sublinear, about as long as propodus; rostrum. Eyes of medium size, ovoid. posterior margin of propodus with 3 setae; dac-

with 7-9 anterior Antenna 1 (fig. 8b) quite setose on peduncle tylus setae on margin. and flagellum; flagellum 7- to 8-segmented, 1 Pereiopod 7 (fig. 9g) poorly setose; merus

shorter than shorter than or 2 aesthetascs on each segment except first, carpus; carpus pro-

and last. with 2 of second podus; propodus groups posterior

Antenna 2 (fig. 8c), flagellum 9- to 10- setae; dactylus armed with 14-16 anterior setae. segmented, heavily setose. Pleopodal peduncles (fig. lOh) each with 5-6

Labrum (fig. 8d) with slightly emarginate retinacula; exopodite of first pleopod 16- free margin. segmented.

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8. Haustorioides 6.7 from head Fig. nesogenes n. sp. (￿ holotype mm Myeongsasimri, Sinjido): a, (scale AB); b, antenna

1 antenna2 labrum mandibles labium maxilla 1 maxilla 2 (AB); c, (AB); d, (AD); e, (AD); f, (AD); g, (AD); h, (AD); i, maxilliped (AC).

Epimeral plate 1 (fig. 9h) with blunt (two of which short and stout). posterodistal corner; plates 2 and 3 with Apex of uropod 3 (fig. 10k) with 2-3 setae; medium-small point. length of longest seta about half that of

Uropod 1 (fig. lOi), peduncle with 5-6 peduncle.

spines. Telson slightly longer than wide, cleft very

Uropod 2 (fig. 10j) with 4 peduncular spines shallow.

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9. Haustorioides 6.7 mm from Sinjido): 1 b, Fig. nesogenes n. sp. (￿ holotype Myeongsasimri, a, gnathopod (scale AB); gnathopod 2 (AB); c, pereiopod 3 (AB); d, pereiopod 4 (AB); e, pereiopod 5 (AB); f, pereiopod 6 (AB); g, pereiopod

7 (AB); h, epimeral plates 1 to 3 (AC).

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10. Haustorioides 9.2 mm from mm from Fig. nesogenes n. sp. (a, c-g, m, ￿ allotype Myeongsasimri, Sinjido; b, ￿ 3.3

Simok, Dochodo; h-k, ￿ holotype 6.7 mm from Myeongsasimri, Sinjido; 1, n, ￿ 4.1 mm from Simok, Dochodo): a, gnathopod 1 ￿ (scale AC); b, gnathopod 1 ￿ (AF,); c, gnathopod 2 ￿ (AC); d, oostegite of gnathopod 2 ￿ (AB); e,

of of oostegite of pereiopod 3 ￿ (AB); f, oostegite pereiopod 4 ￿ (AB); g, oostegite pereiopod 5 ￿ (AB); h, pleopod

1 ￿ (AD); i, uropod 1 ￿ (AD); j, uropod 2 ￿ (AD); k, uropod 3 ￿ (AF); 1, uropod 3 ￿ (AF); m, telson ￿ (AF); n, telson ￿ (AF).

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1 in the but less in Dochodo Female: Coxal plate (fig. 10a) anterodistally type-locality so (fig.

sloping. 10b) and Bigumdo.

Gnathopods 1 and 2 (figs. 10a, c) more setose Etymology. — The proposed specific name than in male.

nesogenes is derived from the Greek, meaning Gnathopod 1, anterior margin of basis with 4 "archipelago" (=Dadohae in Korean) where longer setae and 7 small distal setae; basal part the animal is found. of lobe line of 7 of carpal with setae; tip dactylus

with 2 minute Distribution and — The blades, unguis present. ecology. species

anterior of found in the southwestern islands of the Gnathopod 2 (fig. 10c), margin was

basis with 7-8 distal setae; propodus and dac- Korean peninsula (fig. 1).

in 1 in and this the tylus as gnathopod shape size; carpal Ecologically species occupies upper

lobe with of 7 basal the tidal In row setae. part of zone. Bigumdo (maximum

above the Oostegites on gnathopod 2 and P5 (figs. lOd, tidal range ca. 4.0 m), it was collected

those tide with H. while g) subtriangular, distally narrowing; on mean level together koreanus,

P3 and P4 (figs. lOe, f) distally widened. H. indivisus occupies the whole tidal range of the

of sandy beach. The same zonal distribution the

Remarks. — the in Dochodo. By hardly emarginate species was observed

In females found telson, the present species differs from all other May 1986, ovigerous were

Korean Haustorioides species. in the type-locality. The allotype (body length

From H. koreanus the be distin- 111 size 0.31 species can 9.2 mm) carried eggs, egg 0.26 x

guished by the poor setation on the posterior mm.

of margin P7, the small numberof spines on the

of Haustorioides indivisus n. sp. peduncle uropod 1, the long apical seta on

uropod 3, the equal acumination of the (Figs. 11-13)

corner of 2 and posterodistal epimeral plates 3, Material examined. —Prov. Kyeongki, Yongyoodo, the of anterior the presence marginal setae on Eulwangri; 27 Apr. 1986, 1 O" holotype (ZMA Amph. basis of and the lack of addi- 108.335a), 1 ovigerous 9 allotype (ZMA Amph. gnathopod 1, an

108.335b), 8 99 (6 ovigerous) and 7 CO* paratypes (ZMA tional row of submarginal setae on maxillipedal and 7 and 7 Amph. 108.335c), ovigerous 99 CTC para- palp segment 2. types (IMS). From H. and H. indivisus this latipalpus Prov. Chungnam, Seosan, Hakampo; 19 Oct. 1986, species differs in the dense setation of the anten- 4 crcr and 2 99.

Prov. Chungnam, Malipo; 21 Oct. 1986, 1 OV nae and of the dactylus of P6-P7, the numerous Prov. Chungnam, 9 2 CO" and slender terminal the of the Daecheon; May 1986, setae on dactylus 5 ovigerous 99- and the of maxilliped, presence an apical seta Prov. Cheonbuk, Okgu, Osikdo; 24 July 1986, 5 juvs on the of maxilla The on P5 palp 1. oostegites Prov. Cheonnam, Bigumdo, Gurimri; 19 Oct. 1986, resemble those of H. indivisus but differ from H. 380 specimens. latipalpus. Description. — A small species (adult male Variations are found between the popula- 5.9-6.3 female 6.3-6.5 mm, ovigerous mm). tions of this species. Preadults (body length 3.3-

4.6 from Dochodo and have mm) Bigumdo a Male: Lateral cephalic lobes (fig. 11a) slightly deeper telson cleft (about 1/16 of telson rounded triangular, not reaching end of length, fig. lOn) and a single apical seta on rostrum. Eyes medium small, ovoid.

3 while adults from the Antenna 1 uropod (fig. 10-1), type- (fig. lib), flagellum 7-segmented,

show cleft telson each locality, Sinjido, a barely (fig. with relatively short, scarce setae; segment

and 3 with 1 aesthetascs first and last 10m) 2-3 apical setae on uropod (fig. or 2 except one.

in Antenna 2 slender 10k). Another variation can be seen the (fig. 11c) with peduncle;

of female 1 8- each shape the coxal plate (fig. 10a) flagellum to 9-segmented, segment which is more strongly pointed posterodistally distally with 2 long and some short setae.

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11. Haustorioides indivisus 5.9 from head Fig. n. sp. (￿ holotype mm Eulwangri, Yongyoodo): a, (scale AB); b, antenna

1 2 labrum mandibles labium maxilla 1 maxilla 2 (AB); c, antenna (AB); d, (AC); e, (AC); f, (AC); g, (AC); h, (AC);

i, maxilliped (AC); j, uropod 1 (AB); k, uropod 2 (AB); 1, uropod 3 (AC); m, telson (AC).

Labrum (fig. lid) emarginate at free Labium (fig. llf) of the usual structure; left

lobe without margin. outer groove.

Mandible (fig. lie): Right lacinia mobilis Palp of maxilla 1 (fig. llg) 1-segmented,

with 3-4 medium-sized and 4-5 fine teeth; left without apical seta. Inner lobe short, with two

lacinia mobilis with 4 robust teeth. setae plumose on apex.

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2 11 of the usual Telson Maxilla (fig. h) structure; (fig. 11m) never cleft; tapering

inner margin of inner lobe with 3 strong distally; slightly longer than wide.

plumose setae. Female: Gnathopod 1 (fig. 13a), basis with 3 of Maxilliped (fig. lli), arrangement 3 den- medioproximal and 4 anterodistal setae; basal inner lobe inner tiform spines on irregular; part of carpal lobe with row of 5 setae; dactylus margin with 4 plumose setae. Outer lobe with unguis and 3 additional blades. medium short, not extending to outer tip of Gnathopod 2 (fig. 13b), anterior margin of Inner of palp segment 1. margin palp segment basis with 2 distal setae; shape and size of pro- somewhat of the 2 convex; dactylus linear; one podus and dactylus similar to those in apical spines on dactylus thicker and serrate. gnathopod 1; carpal lobe with 4 basal setules. Coxal plates 1-4 (figs. 12a-d) sparsely setose Oostegites on gnathopod 2 and pereiopod 5 on lower margins. (figs. 13b, e) subtriangular, distally narrowing. Gnathopod 1 (fig. 12a), basis with 3 setae

medioproximally, and 3 small setae antero- Remarks. — From other Korean species of

lobe with 2 small basal Haustorioides this distin- distally; carpal setules; species can be easily

palmar angle of propodus with 2 slender, bifid guished by its entire and distally tapering

spine-teeth; dactylus short, stout, without telson. The species resembles H. latipalpus and

these for unguis. H. nesogenes, see remarks under taxa

Gnathopod 2 (fig. 12b), carpal lobe with their morphological differences.

single basal setule only; length of propodus

— The indivisus Etymology. specific name about 1.8 times that of gnathopod 1. (from the Latin, meaning "not cleft"), alludes Pereiopods 3 and 4 (figs. 12c, d), posterior to the uncleft telson of the animal. margin of merus and carpus relatively poorly

setose. Distribution and ecology. — The species

5 is distributed Pereiopod (fig. 12e) moderately setose; widely throughout the west coast

merus wider than of of Korea For its zonal distribution long; posterior margin pro- (fig. 1). on

podus with single seta; dactylus with 0-1 intertidal sandy beaches, see the remarks under

anterior seta. H. nesogenes.

6 less found in 1986 Pereiopod (fig. 12f) setose; carpus Ovigerous females were April

linear, shorter than propodus; posterior margin in the type-locality (Yongyoodo) and in May of carpus and propodus armed with 1-2 setae; 1986 in Daecheon, with a small number of

with 3-4 The dactylus setae anteriorly. eggs. allotype (body length 6.3 mm) car-

7 ried 21 size 0.25 x 0.32 Pereiopod (fig. 12g) sparsely setose; carpus eggs, egg mm.

shorter than of propodus; posterior margin pro-

podus armed with single bundle of setae; dac- GENERAL REMARKS

tylus with 7-8 anterior setae.

Pleopodal peduncles (fig. 12h) each with 5-6 Outside Korea, Haustorioides japonicus Kami-

of first 1977 be related retinacula; exopodite pleopod 15- to 16- hira, seems to very closely to the

segmented. three small Korean species of Haustorioides (H.

Epimeral plate 1 (fig. 12i) with medium- nesogenes, H. indivisus, and H. latipalpus).

small posterodistal point; plates 2 and 3 with Similarities between the three Korean species

and acute medium-large point. H. japonicus are: Equal length of

Uropod 1 (fig. llj) with 3-4 short, slender posterodistal point on epimeral plates 2 and 3;

small but distinct lobe of spines on peduncle. inner labium;

with 3 and intervals of dentiform Uropod 2 (fig. Ilk) stout 1 small irregular three spines on

peduncular spines only. inner lobes of maxilliped (two innerones closely

Apex of uropod 3 (fig. 11-1) with 1-2 setae; together and another one apart from them);

of 3. length of setae about half that peduncle. relatively long apical seta on uropod

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12. Haustorioides 5.9 from Fig. indivisus n. sp. (￿ holotype mm Eulwangri, Yongyoodo): a, gnathopod 1; b, gnathopod

2; 3; d, 1 c, pereiopod pereiopod 4; e, pereiopod 5; f, pereiopod 6; g, pereiopod 7; h, pleopod 1; i, epimeral plates to 3. Scales: all AB.

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13. Haustorioides indivisus n. 6.3 from b, Fig. sp. (￿ allotype mm Eulwangri, Yongyoodo): a, gnathopod 1; gnathopod

2 with oostegite; c, oostegite of pereiopod 3; d, oostegite of pereiopod 4; e, oostegite of pereiopod 5. Scales: all AB.

In his original description, Kamihira (1977) with long spines only, and densely setose dac-

wrote that “H. japonicus does not have inner tylus of P5-P7 (5-7, 20-25, and 40-50 setae, lobes of labium and has proboscoid epistome." respectively).

distinct inner have But a topotype has lobes of the Bousfield & Tzvetkova (1982) separated labium and a normal He Eohaustorioides from the epistome. figured japonicus genus three regularly implanted dentiform spines on Haustorioides. The criteria on which the new the inner lobes of established the maxilliped, but a topotype genus was are not clear. However, shows irregular intervals of the dentiform if grounds exist in the shape of the telson and in

in the three small of Hausto- the of this spines as species strong process epimeral plate 2, must rioides. be reconsidered, because the cleft of the telson

However, several marked differences are is quite variable as seen in the three Korean

in H. Uncleft but three have present japonicus: not distally species of Haustorioides (all species tapering telson, peduncle of uropods 2 and 3 equal size of epimeral corners 2 and 3, but H.

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indivisus has uncleft H. has 1972. The marine fauna of New Zealand: Algae- an telson, nesogenes a , living littoral (Crustacea Amphipoda). H. cleft hardly cleft telson, and latipalpus has a Bull. N.Z. Dep. scient. ind. Res., 210 (=Mem. telson). N.Z. oceanogr. Inst., 62): 1-216.

BOUSFIELD, E. L., 1970. Adaptive radiation in sand-

burrowing amphipod . Chesapeake Sci., ACKNOWLEDGEMENTS 11 (3): 143-154.

I would like thanks Prof. Dr. 1981. Evolution in North coastal marine to express my special to J. , Pacific

H. Stock of the of Taxonomic Institute Zoology, Univer- amphipod crustaceans. In: G. G. E. SCUDDER & J. L.

sity of Amsterdam (ITZ) for giving me anopportunity for REVEAL eds., Evolution today, Proceedings of the

this study, for his practical advice, and for reading the second International Congress of Systematic and

manuscript. Drs. E. L. Bousfield of the National Museum Evolutionary Biology, Univ. British Colombia, Van-

of Canada, Ottawa and Sung Yun Hong of the Institute couver, 17-24 July 1980: 69-89 (Hunt Institute for

of Marine National Sciences, Fisheries University of Botanical Documentation & Carnegie-Mellon

Pusan (IMS, NFUP) are gratefully acknowledged for their University, Pittsburgh).

critical ofthe and for their continuous 1982. Gammaridea. In: S. P. PARKER reading manuscript , Amphipoda:

interest in this work. Drs. Pyung Chin and Yong Joo ed., Synopsis and classification of living organisms,

Kang of the NFUP, Pusan are thanked for encouraging 2: 254-285 (McGraw-Hill, New York).

E. Studies me during the fieldwork. I wish to thank Dr. S. Pinkster, BOUSFIELD, L. & N. L. TZVETKOVA, 1982. on

Ir. J. Notenboom, and Mr. D. Platvoet of the ITZ, Dogielinotidae (Amphipoda-Talitroidea) from the

for their the shallow waters of the North Pacific Amsterdam, helpful comments on manu- region. Explora-

I indebted script. am to Dr. Y. Kamihira of Hakodate tions of the Fauna ofthe Seas, 29 (37): 76-94 [in Rus-

College, Hokkaido, Japan for supplying topotypic sian with English summary].

material of Haustorioides japonicus. My special thanks are GURJANOVA, E., 1962. Bokoplavy severnoi chasti Tixogo

due to Mr. Chae Woo Ma ofthe IMS, NFUP, Pusan for Okeana (Amphipoda-Gammaridea)chast' 1. Opred.

his assistance with the fieldwork. Faune S.S.S.R., 74: 1-440 [in Russian],

This macrobenthos study has been financially supported by the Jo, Y. W., 1983. Intertidal zonation of on

Netherlands Universities Foundation for International Dadaepo sandy beach, Korea. M. Sc. thesis, Nat.

Cooperation (NUFFIC). Fish. Univ. Pusan: 1-35 [in Korean with English

summary].

KAMIHIRA, Y., 1977. A new species of sand-burrowing

marine amphipods from Hokkaido, Japan. Bull. REFERENCES Fac. Fish. Hokkaido Univ., 28 (1): 1-5.

marine S. BARNARD, J. L., 1967. New and old dogielinotid KIM, H. & C. B. KIM, 1987. Marine gammaridean

Amphipoda. Crustaceana, 13: 281-291. Amphipoda(Crustacea) of Cheju Island and its adja-

1969. The families and of marine cent waters, Korea. Korean J. Zool., 3: 1-23. , genera gamma- syst.

ridean Amphipoda. Bull. U.S. natn. Mus., 271: OLDEVIG, H., 1958. On a new aberrant talitrid from the

1-535. Island of Sachalin. Arkiv Zool., 11: 343-347.

Received: 18 December 1987

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