the caprellidea (malacostraca: amphipoda) from mirs … 2014/hong.pdf · caprella hirayamai, n....

THE CAPRELLIDEA (MALACOSTRACA: AMPHIPODA) FROM

MIRS BAY, HONG KONG, WITH THE DESCRIPTION OF A NEW GENUS

AND TWO NEW SPECIES

J. M. Guerra-Garcıa and I. Takeuchi

(JMGG, correspondence) Laboratorio de Biologıa Marina, Departamento de Fisiologıa y Biologıa Animal,

Facultad de Biologıa, Universidad de Sevilla, Apdo. 1095, E-41080, Sevilla, Spain (e-mail: [email protected]);

(IT) Department of Life Environment Conservation, Faculty of Agriculture, Ehime University, 3-5-7 Tarumi,

Matsuyama, Ehime 790-8566, Japan (e-mail: [email protected])

A B S T R A C T

The taxonomic study on the Caprellidea collected from the shallow waters of north-west area of Mirs

Bay, Hong Kong revealed the distribution of four species in this area. Of them, Tropicaprella minuta, new

genus, new species, and Caprella hirayamai, new species, are described. Tropicaprella, new genus, is

close to Heterocaprella Arimoto, 1976, but differs by the absence of the clavate projection on the

pereonite 4, the elongate pereopods 3 and 4 and the five-articulate pereopod 5. The closest species to

Caprella hirayamai, n. sp., are Caprella californica Stimpson, 1857, and Caprella scaura Templeton,

1836, but C. hirayamai can be easily distinguished by less elongated pereonite 1, short basis of gnathopod2 and oval propodus with dorsal setae in gnathopod 2.

Since the publication of Mayer’s monographsof the Caprellidea of the world (Mayer, 1890,1903), no systematic studies on the Caprellideafrom the Chinese coasts have been carried out.According to McCain and Steinberg (1970), thefollowing nine species of the Caprellidea werereported from Chinese coasts: Hemiaeginaminuta Mayer, 1890; Caprella acanthogasterMayer, 1890; C. bacillus Mayer, 1903; C.bispinosa Mayer, 1890; C. californica Stim-pson, 1856; C. drepanochir Mayer, 1890; C.equilibra Say, 1818; C. penantis Leach, 1814;and C. rhopalochir Mayer, 1890. In addition,several species of Caprella have been reportedin scattered ecological studies (e.g., Brawleyand Fei, 1987, 1988; Huang et al., 1992).Especially, Brawley and Fei (1987, 1988)reported the presence of five species of Caprellafrom the northern coast of China: C. equilibraSay, 1818; C. irregularis Mayer, 1890; C.kroyeri de Haan, 1849; C. scaura Templeton,1836; and C. septentrionalis Kroyer, 1838.

Recently, the necessity for conserving marinehabitats along the Hong Kong coast has beenreflected by the designation in 1996 of HongKong’s first Marine Parks and Reserves. Twoparks and one reserve received designation on 5July 1996 (Morton, 2000). These areas possesa wide diversity of marine intertidal and subtidalhabitats and high richness of corals, mangroves,seagrasses, dynamic sand beaches, and lagoons.The amphipods were collected from various

marine habitats to 24 meters depth from theTolo Channel area of northeastern New Terri-tories, Hong Kong, including the two marineparks, Yan Chau Tong Park and Hoy Ha WanPark, during the Second International MarineBiological Workshop in April 1984. Of thecollections, Hirayama (1986a, b; 1991) pub-lished a series on the taxonomy of theGammaridea, including twelve species of Co-rophium, Corophiidae (Hirayama, 1986a), fourspecies of Dexaminiidae (Hirayama, 1986b),and six species of Ampeliscidae (Hirayama,1991). Based on this collection, Hirayama(1986a) estimated that the gammaridean amphi-pod fauna of Hong Kong and its adjacentregions is less diverse than other waters of theWest Pacific in the Northern Hemisphere.

In the present study, four species of theCaprellidea belonging to Hirayama’s collectionfrom Hong Kong, and containing one newgenus and two new species, are reported.Taking into account that phylogeny and higherclassification of the Caprellidea is still underdebate (e.g., Laubitz, 1993; Takeuchi, 1993;Larsen, 1997), the genera have been groupedconsidering the smaller number of families,following Takeuchi (1993).

MATERIALS AND METHODS

The amphipod material was collected with a 0.25 m2 VanVeen grab, a small-size beam trawl, or direct handlingwithoutgears, from 4 to 24 April 1986 from Tolo Haubour, Tolo and

154

JOURNAL OF CRUSTACEAN BIOLOGY, 23(1): 154–168, 2003

pdi

Texto escrito a máquina

northwest Mirs Bay of northeastern New Territories, Hong

Kong (1148209N, 228309E). A map and details of the stations

were included in Hirayama (1986a). Additional specimens

were collected frommetal panelswhich had been immersed in

Tai TamBay, south of HongKong Island, by Prof. B.Morton.

The specimenswere fixed in 70% ethanol after preservation in

5% Formalin after the collection. Several specimens were

dissected under a stereomicroscope, and permanent mounts

were made in polyvinyl-lactophenol. All figures have been

drawn with the aid of a camera lucida. The type specimens

are deposited in the Australian Museum, Sydney.

SYSTEMATICS

Family Phtisicidae Vassilenko, 1968

Genus Metaproto (Haswell, 1880)Metaproto sp.

Fig. 1

Material Examined.—1 male, 1 female, st. 4, 16 m deep, 4April 1986, Van Veen grab (1148209N, 228309E).

Remarks.—The genus Metaproto comprises

Fig. 1. Metaproto sp. Lateral view. A, male; B, female. Scale bar: 1 mm.

155GUERRA-GARCIA AND TAKEUCHI: CAPRELLIDEA FROM HONG KONG

pdi


Erratum: Coordinates 22º30'N 114º20'E instead of 114º20'N, 22º30'E

pdi


pdi


pdi


only one species, Metaproto novaehollandiae,which was described by Haswell (Haswell,1880). The type material of M. novaehollan-diae, although thought to be deposited in theAustralian Museum, Sydney (McCain andSteinberg, 1970), was lost (Springthorpe andLowry, 1994). Mayer (1903) redescribed thespecies based on the specimens collected fromthe Banda Sea, West of Papua-New Guinea, andLaubitz (1991) figured one specimen in poorcondition from Phillipines. Currently, Metapro-to novaehollandiae from New South Wales,Australia, is undergoing redescription basedon specimens collected near the type locality(Takeuchi, in prep.). The present specimens,collected from Hong Kong, clearly belong to thegenus Metaproto on the basis of the presence ofa five-articulate pereopod 5 and abdomen withonly one pair of appendages. However, theshape of gnathopod 2 and the number ofgrasping spines on the propodus of pereopods3 and 4 in males (three in M. novaehollandiaeand one in the present specimen) indicate thatthese specimens collected from Hong Kongshould be a different species from M. novae-hollandiae. However, only two specimens werefound, and the male is not completely adult,preventing us from assigning these specimens toa new species in the present study.

Family Caprellidae Leach, 1814

Tropicaprella, new genus

Diagnosis.—Flagellum of antenna 2 two-artic-ulate, pereopods 3 and 4 uniarticulate. Pereopod5 five-articulate (although articles 4 and 5 arenot clearly separated). Pereopods 6 and 7elongate, six-articulate. Gills on pereonites 3and 4. Mandibular palp absent. Molar present.Male abdomen with pair of appendages and pairof lobes; female abdomen without appendages.

Type Species.—Tropicaprella minuta, n. sp.

Gender.—Feminine.

Etymology.—The name Tropicaprella was cho-sen because the species was found very close toTropic of Cancer.

Remarks.—Tropicaprella, new genus, is mostclosely related to the genus HeterocaprellaArimoto in flagellum of antenna 2 two-articulate, pereopods 3 and 4 uniarticulate,gills on pereonites 3 and 4, and mandibularpalp absent, while molar present. The genus

Heterocaprella is, so far, composed of twospecies, Heterocaprella clavigera Arimoto,1976, described based on specimens collectedfrom the Korean Strait (Arimoto, 1976a), andHerocaprella krishnaensis Swarupa and Rad-hakrishna, 1983, collected from Indian waters(Swarupa and Radhakrishna, 1983). Tropicap-rella can be distinguished from Herocaprellaby the following characters: (1) males ofHeterocaprella posses a striking clavate pro-jection of pereonite 4, carrying the gills andpereopods at its distal end—this projection,unique in the Caprellidea, is lacking in malesof the new genus Tropicaprella; (2) thepereopods 3 and 4 are two-articulate inHeterocaprella and uniarticulate in Tropicap-rella; (3) the pereopod 5 is 3–4-articulate inHerocaprella and 5-articulate in Tropicaprella.The two species of Heterocaprella showabdominal specific variations, which are usu-ally considered as generic characters in thegroup (see McCain, 1968; Takeuchi, 1993);males and females of Heterocaprella clavigerahave a pair of appendages on the abdomen(Arimoto, 1976a), whereas these appendagesare absent in the species H. krishnaensis(Swarupa and Radhakrishna, 1983).

Tropicaprella minuta, new speciesFigs 2–6

Type Material.—Holotype male (AM P61575), st. 19, 17 mdeep, 4 April 1986, Van Veen grab (1148209N, 228309E ).Allotype female (AM P61576), st. 25, 18 m deep, 8 April1986, Van Veen grab, Mirs Bay outer Tolo Channel(1148209N, 228309E). Paratypes (AM P61577): 2 females,st. 17, 18 m deep, 9 April 1986, on crinoids, Port Islands; 1male, st. 18, 17.5 m deep, 8 April 1986, Van Veen grab(1148209N, 228309E); 1 female, st. 20, 17 m deep, 8 April1986, Van Veen grab (1148209N, 228309E); 6 females, st. 23,18 m deep, 8 April 1986, Van Veen grab, Mirs Bay outerTolo Channel; 1 male, st. 24, 18 m deep, 8 April 1986, VanVeen grab, Mirs Bay outer Tolo Channel.

Description.—Holotype male (AM P61575).Body length 3.2 mm.

Lateral view: Body smooth provided withfew dorsal setulae. Head rounded. Pereonites 2–5 subequal in length; pereonite 7 the shortest.

Gills elongated, length about 2.5 times width.Antenna 1 about 1/4 of body length; pedu-

ncular article 1 subequal to article 3 in lengthbut wider; flagellum 5-articulate. Antenna 2 aslong as antenna 1; swimming setae absent; basalarticle of peduncle with acute projection.

Mouthparts remarkably tiny (about 0.05 mmlength). Upper lip symmetrically bilobed,

156 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 23, NO. 1, 2003

smooth. Mandibular process present; left man-dible with incisor 5-toothed, lacinia mobilis 5-toothed followed by row of three pectinatesetae; right mandible with incisor 5-toothed,lacinia mobilis not toothed (or minutely serrat-ed), followed by two pectinate setae; molarflake absent. Inner lobes of lower lip round:inner and outer lobes smooth apically. Maxilla 1

outer lobe with 7 serrated spines apically; article2 of palp with 4 marginal setae and 2lateral ones. Maxilla 2 inner lobe with 6 apicalsetae, outer lobe with 10 setae. Inner plateof maxilliped rectangular, with 4 setae ondistal end; outer plate with one apical setaeand 2 lateral ones; palp 4-articulate, scarcelysetose.

Fig. 2. Tropicaprella minuta, n. g., n. sp. Lateral view. A, male; B, female. Scale bar: 1 mm.


Fig. 3. Tropicaprella minuta, n. g., n. sp. Male. A, Upper lip; B, Lower lip; C, Maxilliped; D, right mandible; E, leftmandible; F, maxilla 1; G, maxilla 2. Scale bars: 0.05 mm.


Fig. 4. Tropicaprella minuta, n. g., n. sp. Male. A, antenna 1; B, antenna 2; C, gnathopod 1; D, gnathopod 2. Scale bars:0.2 mm.


Gnathopod 1 unique in the Caprellidea;carpus with two strong spines; palm ofpropodus without proximal grasping spines,with small, acute projection proximally, fol-lowed by rounded expansion, minutely serrateand provided with 7 setae, and small projectiondistally; dactylus elongate, strongly divideddistally. Gnathopod 2 inserted near distal endof pereonite 2; basis about 1/3 as long aspereonite 2; propodus elongated, length about 3times width; palm with proximal projectionbearing robust spine and another two smallprojections distally.

Pereopods 3 and 4 uniarticulate, small, lengthabout 1/4 of gills; pereopod 3 with 4 distalsetae, pereopod 4 with one distal seta. Pereopod5, inserted at middle of pereonite 5, five-articulate, although articles 4 and 5 notcompletely separated. Pereopods 6 (missing inmale holotype) and 7 elongate, similar in chara-cteristics but increasing in length respectively;basis to propodus setose, dactylus long, curved;propodus palm without grasping spines.

Abdomen with pair of appendages uniarticu-late, short, rounded, with pair of lateral lobesand single dorsal lobe. Penes elongate.

Fig. 5. Tropicaprella minuta, n. g., n. sp. Male. A, pereopod 3; B, pereopod 4; C, pereopod 5; D, pereopod 6; E, pereopod7. Scale bars: A–C: 0.1 mm; D, E: 0.5 mm.


Allotype female.—(AM P61576).—Body length2.4 mm. Gnathopod 2 inserted on anterior halfof pereonite 2, similar to male gnathopod 2although propodus slightly elongated. Broodlamellae of pereonite 3 and 4 setose. Gills moreelongate than in male. Pereopod 5 inserted onanterior half of pereonite 5. Abdomen withoutappendages, with pair of lateral lobes and dorsallobe.

Etymology.—The specific name ‘‘minuta’’ refersto the small size of the new species.

Remarks.—The tiny new species, althoughcloser morphologically to Heterocaprella spe-cies, superficially resembles Liropus minimusMayer, 1890, especially in size and shape of thebody, but differs easily by the followingcharacters: (1) pereopod 5 is 2-articulate inLiropus minimus and 5-articulate in Tropicap-rella minuta; (2) Liropus minimus has a 3-articulate mandibular palp, and the palp isabsent in Tropicaprella minuta; (3) the shapeof gnathopod 1 and the length and character-istics of pereopods 6 and 7 differ between thespecies.Tropicaprella minuta, n. sp., possesses ex-

tremely long pereopods 6 and 7. They could beuseful in some way as an adaptation to living onsediments.

Genus Caprella Lamarck, 1801Caprella scaura Templeton, 1836

Fig. 7

Caprella scaura Templeton, 1836: 191, 192, pl. 20, fig. 6;

Mayer, 1890: 70–73, pl. 4, figs. 40–51, pl. 6, fig. 41, pl. 7,

figs. 2, 35, 36; Mayer, 1903: 117–120, pl. 5, figs. 13–18,

pl. 10, fig. 11; McCain, 1968: 40–44, figs. 17, 18, 55.Caprella nodosa Templeton 1836: 192–194, pl. 21, fig. 7.Caprella cornuta Dana, 1853: 816, 817.Caprella attenuata Dana, 1853: 817–819.

A more extensive list of synonymies for this species isincluded in McCain and Steinberg (1970).

Material Examined.—1 male, st. 7, 16 m deep, 8 April 1986,Northern coast of Kat Ochan (1148209N, 228309E); 2 malesand 1 premature female, st. 8, 15 m deep, 5 April 1986, algalbed, Kai Kung Tau (1148209N, 228309E); 1 male, st. 9, 14 mdeep, April 1986, algal bed, Kai Kung Tau (1148209N,228309E); 1 female, st. 11, 14 m deep, 9 April 1986, benthictrawl.

Remarks.—Caprella scaura was reported to bedistributed widely among pier pilings in thesame area as the present collection prior to thepresent study (Huang et al., 1992). This speciesis distributed widely along temperate regions allover the world (McCain, 1968; McCain andSteinberg, 1970). Caprella scaura was firstdescribed by Templeton (1836) based onmaterial from Mauritius, South Indian Ocean.Seven varieties have been described by Mayer

Fig. 6. Tropicaprella minuta, n. g., n. sp. A, male abdomen (ventral view); B, female abdomen (ventral view). Scale bar:0.1 mm.


(1890, 1903) and Utinomi (1947). Of thesevarieties, McCain (1968) re-established C.californica Stimpson, 1857, and remanded theothers as junior synonyms of Caprella scauraTempleton. The specimens examined in thepresent study are in good agreement with thefigures and descriptions of Caprella scaura

typica Mayer, 1890 from Brazil (Mayer, 1890;Arimoto, 1976b) and of Caprella scaura ofMcCain (1968) from the Western North Atlan-tic. This species shows a wide variation ofsubstrates, indicating no specific selection of thehabitats. Caprella scaura has been frequentlyrecorded in association with the bryozoan

Fig. 7. Caprella scaura Templeton, 1836. Lateral view. A, male; B, female. Scale bar: 1 mm.


Bugula neritina and other erect bryozoans of thegenus Scrupocellaria, with the seaweeds Sar-gasum spp. and Cystoseira spp., and with theseagrasses Halodule uninervis and Halophilaovalis (see Lim and Alexander, 1986). Takeuchiand Hino (1997) found the species attaching tothe seagrasses Zostera marina and Z. caules-cens, and Serejo (1998) reported the presence ofthe species clinging to the sponge Dysideafragilis. In Coquimbo, the species has been

found living on Bugula neritina and theseaweeds Polysiphonia spp. and Gracilariaspp. (Guerra-Garcıa and Thiel, 2001). Caprellascaura is one of the caprellid species for whichthe behaviour is well described (Lim andAlexander, 1986; Aoki, 1999; Schultz andAlexander, 2001).

Caprella hirayamai, new speciesFigs. 8–11

Fig. 8. Caprella hirayamai, n. sp. Lateral view. A, male; B, female. Scale bar: 1 mm.


Fig. 9. Caprella hirayamai, n. sp. Male. A, Upper lip; B, Lower lip; C, Maxilliped; D, right mandible; E, left mandible;F, maxilla 2; G, maxilla 1. Scale bars: 0.2 mm.


Fig. 10. Caprella hirayamai, n. sp. A–D, male. A, antenna 1; B, antenna 2; C, gnathopod 1; D, gnathopod 2. E, femalegnathopod 2. Scale bars: A, B, D, E: 1 mm; C: 0.5 mm.


Type Material.—Holotype male (AM P61578), st. 1, 20 mdeep, 4 April 1986, Van Veen grab (1148209N, 228309E).Allotype female (AM P61579), st. 13, 14 m deep, 9 April1986, on crinoids, Port Island (1148209N, 228309E). Para-types (AM P61580): 4 juveniles collected together with theallotype female in the same sample; 5 males, 2 prematurefemales, 1 immature female, st. 2, 24 m deep, 4 April 1986,Van Veen grab (1148209N, 228309E); 8 females, 1 immaturefemale, 4 juveniles, st. 5, 16 m deep, 9 April 1986, on thecrinoid Tropiometra afra (1148209N, 228309E); 1 female, st.

6, 15 m deep, 5 April 1986, on algae (1148209N, 228309E); 1female, st. 16, collected from fouling community on metalpanels at Tai Tam.

Diagnosis.—Head with well-developed acuteprojection. Body dorsally smooth except forpair of small tubercles on pereonite 6 in malesand on pereonites 6 and 7 in females; femalesalso provided with dorsal humps on pereonites

Fig. 11. Caprella hirayamai, n. sp. A–D, male. A, pereopod 5; B, pereopod 6; C, pereopod 7; D, abdomen. E, femaleabdomen. Scale bars: A–C: 0.5 mm; D, E: 0.2 mm.


2–5. Basis of gnathopod 2 with distolateralprojection in males; small dense setulae dorsallyon distal half of male propodus.

Description.—Holotype male (AM P61578).Body length 9.4 mm.Lateral view: Head with well-developed

acute projection. Body dorsally smooth exceptfor pair of small tubercles on pereonite 6.Pereonites 3–5 subequal, pereonite 6 thelongest.Gills elongated, length about 3 times width.Antenna 1 longer than half of body; pedun-

cular article 1 subequal to article 3; flagellumbroken distally. Antenna 2 as long as antenna 1peduncle; swimming setae present.Upper lip symmetrically bilobed, slighly

pubescent apically. Mandibular process strong;incisor and lacinia mobilis 5-toothed in bothmandibles; left mandible with three pectinatesetae, right mandible with two pectinate setae;molar flake small, rounded, setose. Inner lobesof lower lip round; inner and outer lobespubescent apically. Maxilla 1 outer lobe with7 serrated spines apically; article 2 of palp with9 marginal setae and 15 lateral ones disposedalong surface. Maxilla 2 inner lobe with 20apical setae, outer lobe with 17 setae. Inner plateof maxilliped rectangular, with 9 plumose setaeand 3 nodular setae (like teeth) on distal end;outer plate with 7 nodular setae (like teeth).Gnathopod 1 propodus with two proximal

grasping spines, dactylus minutely serrated.Gnathopod 2 inserted near distal end ofpereonite 2; basis about half of pereonite 2 inlength, with well-developed projection; propo-dus oval, length about 2.5 times width; palmwith projection bearing robust spine, anotherprojection followed by ‘‘U’’ notch distally.Pereopods 5, 6, and 7 increasing in length,

respectively; basis with carina on posteriormargin; palm of propodus with pair of proximalgrasping spines.Abdomen with pair of appendages uniarticu-

late, pair of lateral lobes and single dorsal lobe.Penes triangular, acute distally.

Allotype Female (AM P61579).—Body length8.1 mm. Body with pair of small tubercles onpereonites 6 and 7 and dorsal humps inpereonites 2–5. Pereonites 2–5 subequal inlength. Gnathopod 2 inserted on anterior halfof pereonite 2; inner projection on basis lacking;propodus without ‘‘U’’ notch. Brood lamellae ofpereonite 3 setose; brood lamella on pereonite 4

not setose. Abdomen with pair of lateral lobescarrying 1 single seta and 1 dorsal lobe.

Etymology.—The species is dedicated to Dr.Akira Hirayama for his contribution to theknowledge of Amphipoda from Indo-Pacificareas. He provided us with the caprellid speci-mens for taxonomic studies.

Remarks.—The new species is close to Caprellacalifornica Stimpson, 1857, and C. scauraTempleton, 1836, in the general lateral viewand the marked acute projection on the head.Based on the comparison using the representa-tive descriptions of C. californica (McCain,1968; Arimoto, 1976b) and C. scaura (Mayer,1890; Arimoto, 1976b), the males of Caprellahirayamai, n. sp., can be easily distinguishedfrom these two species mainly by the followingcharacteristics: (1) in C. hirayamai, pereonite 1is not longer than pereonites 3 and 4, while in C.californica and C. scaura, pereonite 1 iselongate, more than twice of pereonite 3 inlength; (2) in C. hirayamai, the basis ofgnathopod 2 is shorter than half of the pereonite2, whereas in C. californica and C. scaura, it isclearly longer than half of the pereonite 2; (3) inC. hirayamai, the propodus of gnathopod 2 isoval and provided with dorsal setulae near distalend, but it is elongate and without dorsal setaein C. scaura, in which it is covered withnumerous fine setae on all its surface; (4) thedistal article of the peduncle of antenna 1 isshort in C. hirayamai and elongate in C.californica and C. scaura.The females of Caprella hirayamai, n. sp.,

strongly resemble females of C. californica andC. scaura. However, the relative larger andwider gills distinguish C. hirayamai from C.californica and C. scaura even in females.Caprella hirayamai, n. sp., has been found fromsediments, on algae, crinoids, and as a part ofthe fouling community during the present study.This indicates that Caprella hirayamai, as wellas C. scaura, shows no substrate specificity,being a common species around Hong Kong,and probably along the southern coast of China.

ACKNOWLEDGEMENTS

We are very grateful to Dr. Akira Hirayama for providingus the specimens of the Caprellidea for the present work.This study was partially supported by a grant ‘‘Formacion deProfesorado Universitario’’ from the Spanish Ministry ofEducation, Culture and Sport to JMGG, and the SpecialResearch Fund from the Ehime University to IT.


LITERATURE CITED

Aoki, M. 1999. Morphological characteristics of young,maternal care behaviour and microhabitat used bycaprellid amphipods.—Journal of the Marine BiologicalAssociation of the United Kingdom 79: 629–638.

Arimoto, I. 1976a. Heterocaprella clavigera n. gen., n. sp.a unique new caprellid with remarkable sexual dimor-phism, from the Korean Straits (Amphipoda, Caprelli-dea).—Crustaceana 30: 43–48.

———. 1976b. Taxonomic studies of caprellids (Crustacea,Amphipoda, Caprellidae) found in the Japanese andadjacent waters.—Special Publications from Seto MarineBiological Laboratory, Kyoto University, Series III: i–v1 1–229.

Brawley, S. H., and Fei, X. G. 1987. Studies of meso-herbivory in aquaria and in an unbarricaded mariculturefarm of the Chinese coast.—Journal of Phycology 23:614–623.

———, and ———. 1988. Ecological studies ofGracilaria asiatica and Gracilaria lemaneiformis inZhanshan Bay, Qingdao.—Chinese Journal of Oceanog-raphy and Limonology 6: 22–36.

Dana, J. D. 1853. Crustacea.—United States ExploringExpeditions 14: 689–1618.

Guerra-Garcıa, J. M., and M. Thiel. 2001. La fauna decaprelidos (Crustacea: Amphipoda: Caprellidea) de lacosta de Coquimbo, centro-norte de Chile, con una clavetaxonomica para la identificacion de las especies.—Revista Chilena de Historia Natural 74: 873–883.

Haswell, W. A. 1880. On some additional new genera andspecies of amphipodous crustaceans.—Proceedings of theLinnean Society of New South Wales 4: 319–350.

Hirayama, A. 1986a. Marine Gammaridean Amphipoda(Crustacea) from Hong Kong I. The family Corophidae,genus Corophium. Pp. 449–485 in B. Morton, ed.Proceedings of the Second International Marine Bi-ological Workshop: The Marine Flora and Fauna ofHong Kong and Southern China. Hong Kong UniversityPress, Hong Kong.

———. 1986b. Marine Gammaridean Amphipoda (Crusta-cea) from Hong Kong II. The family Dexaminidae. Pp.487–501 in B. Morton, ed. Proceedings of the SecondInternational Marine Biological Workshop: The MarineFlora and Fauna of Hong Kong and Southern China.Hong Kong University Press, Hong Kong.

———. 1991. Marine Ampeliscidae (Crustacea: Amphi-poda) from Hong Kong.—Asian Marine Biology 8: 77–93.

Huang, Z. G., S. K. Yan, S. Lin, and D. Q. Zheng. 1992.Biofouling communities on pier piling in Mirs Bay. Pp.529–543 in B. Morton, ed. The Marine Flora and Faunaof Hong Kong and Southern China III. Volume 2. HongKong University Press, Hong Kong.

Larsen, K. 1997. A new species of Metaprotella (Crustacea:Amphipoda: Caprellidea) from east Africa, with key tothe genera of Protellidae and discussion of genericcharacters.—Journal of Natural History 31: 1203–1212.

Laubitz, D. R. 1991. Crustacea Amphipoda Caprellidea:Caprellids from the western Pacific (New Caledonia,Indonesia and the Philippines). Pp. 101–123 in A.Crosnier, ed. Resultats des Campagnes MUSORSTOM.-

Vol. 9. Memoires du Musee National d’Histoire Naturellede Marseille, Paris.

———. 1993. The Caprellidea (Crustacea: Amphipoda):toward a new synthesis.—Journal of Natural History 27:965–976.

Lim, S. T. A., and C. G. Alexander. 1986. Reproductivebehaviour of the caprellid amphipod Caprella scauratypica Mayer, 1890.—Marine Behaviour and Physiology12: 217–230.

Mayer, P. 1890. Die Caprelliden des Golfes von Neapel andder angrenzenden Meeres-Abschnitte.—Fauna and Florades Folfes von Neapel 17: 1–157.

———. 1903. Die Caprellidae der Siboga Expedition.—Siboga-Expeditie 34: 1–160.

McCain, J. C. 1968. The Caprellidea (Crustacea: Amphipo-da) of the Western North Atlantic.—Bulletin of theUnited States National Museum 278: 1–16.

———, and J. E. Steinberg. 1970. Amphipoda I. Caprelli-dea I. Fam. Caprellidae. Pp. 1–78 in H. E. Gruner and L.B. Holthuis, eds. Crustaceorum Catalogus, Pars 2.

Morton, B. 2000. Coastal zone management for marineconservation in Hong Kong: the need for regional co-operation in Southern China. Pp. 3–33 in B. Morton, ed.Proceedings of the Tenth International Marine BiologicalWorkshop: The Marine Flora and Fauna of Hong Kongand Southern China. Hong Kong University Press, HongKong.

Serejo, C. S. 1998. Gammaridean and caprellidean fauna(Crustacea) associated with the sponge Dysidea fragilisJohnston at arraial do Cabo, Rio de Janeiro, Brazil.—Bulletin of Marine Science 63: 363–385.

Springthorpe, R. T., and J. K. Lowry. 1994. Catalogue ofcrustacean type specimens in the Australian Museum:Malacostraca.—Technical Reports of the AustralianMuseum 11: 1–134.

Stimpson, W. 1857. The Crustacea and Echinodermata ofthe Pacific shores of North America.—Boston Journal ofNatural History 6: 444–532.

Schultz, M. K., and C. G. Alexander. 2001. Aggressivebehaviour of Caprella scaura typica Mayer, 1890(Crustacea: Amphipoda).—Marine and Freshwater Be-haviour and Physiology 34: 181–187.

Swarupa, K. M. C., and Y. Radhakrishna. 1983. Metacap-rella krishnaensis n sp. a new caprellid from Indianwaters (Amphipoda, Caprellidea).—Crustaceana 44:54–60.

Takeuchi, I. 1993. Is the Caprellidea a monophyleticgroup?—Journal of Natural History 27: 947–964.

———, and A. Hino. 1997. Community structure ofcaprellid amphipods (Crustacea) on seagrasses in OtsuchiBay, Northeastern Japan, with reference to the associationof Caprella japonica (Schurin) and Phyllospadix iwaten-sis Makino.—Fisheries Science 63: 327–331.

Templeton, R. 1836. Descriptions of some undescribedexotic Crustacea.—Transactions of the EntomologicalSociety of London 1: 185–198.

Utinomi, H. 1947. Caprellidae of Japan and adjacentwaters.—Seibutsu, Supplement (1): 68–82. (In Japanese).

RECEIVED: 1 April 2002.ACCEPTED: 19 June 2002.


the caprellidea (malacostraca: amphipoda) from mirs … 2014/hong.pdf · caprella hirayamai, n....

Documents