The copepod fauna of the Gulf of Carpentaria, and its Indo-West Pacific affinities

561

Netherlands Journal of Sea Research 25 (4): 561-572 (1990)

THE COPEPOD FAUNA OF THE GULF OF CARPENTARIA, AND ITS INDO-WEST PACIFIC AFFINITIES B.H.R. OTHMAN 1,2, J.G. GREENWOOD 2 and P.C. ROTHLISBERG3 1Department of Zoology, Faculty of Life Sciences, Universiti Kebangsaan Malaysia, 43600 Bangi, Selangor, Malaysia 2Department of Zoology, University of Queensland, St. Lucia 4067, Australia 3CSIRO, Division of Fisheries Research, Marine Laboratories, RO. Box 120, Cleveland 4163, Australia

ABSTRACT Copepods from the Gulf of Carpentaria, the tropical, northern Australian waters adjacent to the Arafura Sea, were collected from 23 sites spanning the entire Gulf during 10 cruises in the period August 1975 to May 1977. Samples were taken using plankton nets with mesh size of 142 #m. One hundred and two species of copepods were identified, 68 species belonging to the sub-order Calanoida, 30 to the Cyclopoida and four to the Harpacticoida. Thirteen of these species were new to science; descriptions of seven have so far been published. Ninety five of the species collected in this study are new records for the Gulf region, and 23 are new records for waters of the Australian continent. The faunal composition is characteristically warm water neritic, and shows similarity to that of Southeast Asia, having at least 88 species in common. Comparisons of copepod species records from the Gulf of Carpentaria with those from the northeastern Pacific coast of Australia (Great Barrier Reef and Moreton Bay) show 79 species to be common to these areas. Of all the species collected in the Gulf, three have been previously recorded from the Indian Ocean but not the Pacific, and five are otherwise only known from the Pacific Ocean. The Gulf is regarded as forming the southeasternmost region of the Indo-pacific marine domain, within which most of the 13 new species encountered in this study appear to be endemic. Species distribution patterns are discussed in relation to postglacial and present-day hydrodynamics of the region. 1. INTRODUCTION The waters surrounding Australia belong to both tropical and temperate regions. Correspondingly, the copepod fauna in the north is markedly different from that in southern Australian waters. Regional studies which include copepods have been published for the following marine areas: the south (NICHOLLS, 1944;

ARNOTT, 1974a, 1974b; TAW, 1974, 1975, 1978; KIMMERER & MCKINNON, 1985); the southeast (DAKIN & COLEFAX, 1940; KOTT,1957; TAFE& GRIFFITHS, 1983; TRANTER et al., 1983); the subtropical Moreton Bay region of the central east coast (THWIN, 1972; GREENWOOD, 1976, 1977, 1978a, 1978b, 1979, 1980, 1981, 1982a, 1982b); the northeast, in particular areas within the Great Barrier Reef (RUSSELL & COLEMAN, 1934; FARRAN, 1936, 1949; GRIGG, 1972; ALLDREDGE & KING, 1977; HODGSON, 1982; JACOBY & GREENWOOD, 1988); the northwestern and western waters (MARKINA, 1971, 1972a, 1976; TRANTER, 1977). Few copepod studies have been made in waters of the northern tropical Australian shelf, that relatively large area adjacent to the Timor and the Arafura Seas. The most extensive was that by MARKINA (1972b), who examined 185 samples taken on several cruises in the Arafura and the Timor Seas, the northwest coast, and the Gulf of Carpentaria. Taxonomic works on copepods in samples from the Gulf of Carpentaria have been published on Labidocera (GREENWOOD & OTHMAN, 1979; FLEMINGER et al., 1982), and quantitative zooplankton biomass estimates have been made by MARKINA(1971) and ROTHLISBERG & JACKSON (1982). This lack of detailed knowledge of the species composition and distribution initiated the present study. The material studied stemmed from CSIRO oceanographic cruises in the Gulf of Carpentaria during August 1975 to May 1977 (ROTHLISBERG & JACKSON, 1982). Acknowledgements.--We would like to thank Chris Jackson and Bob Pendrey for their assistance in collecting and curating the plankton samples. R. Othman's participation in this study was supported by a Commonwealth Scholarship from the Australian Government and the Universiti Kebangsaan Malaysia. The sampling programme conducted for the east coast of the Malay Peninsula was supported by Petronas Carigali Sdn. Bhd. Dr. M.A. Baars made many valuable contributions to the preparation of this paper, and Prof. Dr. W. Vervoort kindly advised on some taxonomic aspects, for all of which we are most grateful.

562

B.H.R. OTHMAN, J.G. GREENWOOD & P.C. ROTHLISBERG

2. MATERIAL AND METHODS The Gulf of Carpentaria (Fig. 1), herein referred to as 'the Gulf', is a U-shaped basin lying within latitude 11° to 17°30'S and longitude 136° to 142° E. This partly enclosed body of tropical water has an approximate area of 3.69.105 km 2 with a surrounding coastline of about 1 900 km, excluding shores of islands and bays. Its size is therefore about equal to that of the Gulf of Thailand (Fig. 2). Greatest width of the Gulf is 650 km at latitude 14°45'S and its greatest length is 800 km along longitude 140°E. Maximum depth is about 70 m, with the 20 m and 40 m depth contours approximately paralleling the coastline (Fig. 1). Depths greater than 60 m are confined to a relatively small area (15% of total area) of irregular shape oriented north-south in the northeast part of the Gulf. The Gulf opens northwards into the Arafura Sea which then joins the Timor Sea and hence the Indian Ocean in the west, and the Banda Sea in the north (Fig. 2). The Banda Sea joins the Pacific Ocean northwards through the many islands of Indonesia. The eastern part of the Arafura Sea is connected to the Coral Sea, and hence the Pacific Ocean again, through the narrow (150 km) and shallow (20 m) Torres Strait between Papua New Guinea and Cape York.

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136°

J

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138°

~ I

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140°

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Fig. 1. Gulf of Carpentaria sampling locations used by ROTHLISBERG& JACKSON(1982). Subset of stations used for copepod analysis in this study are indicated by squares. Depth contours in metres.

Pacific Ocean

CerS°L~'~,~, •

'

° '

•

Indian Ocean

Fig. 2. Locations in Indo-west Pacific used for comparison with Gulf of Carpentariafauna. Square denotes the location of sampling by Othman et al. (1987). AS: Arafura Sea; AI: Aru Island; BAI: Buru/Ambon Islands; BS: Banda Sea; CelS: Celebes Sea; CerS: Ceram (Seram) Sea; CS: Coral Sea; Fs: Flores Sea; GOC: Gulf of Carpentaria; GO~. Gulf of Thailand; JS: Java Sea; MSt: Macassar Strait; SaS: Sahul Shelf; SUS: Sunda Shelf; SCS: South China Sea; SS: Sulu Sea; StM: Strait of Malacca; TS: Timor Sea; TSt: Torres Strait. The CSIRO cruises in the Gulf, 10 in total during the period August 1975 to May 1977, covered a grid of 70 stations (Fig. 1). Details of the sampling programme and of sample handling methods are given in ROTHLISBERG & JACKSON (1982). Samples for this study were taken from a subset of 23 stations from all 10 cruises. Plankton nets having 0.25 m 2 mouth area with mesh size of 142 #m were towed in stepped-oblique hauls from the sea surface to the near-bottom at one or two intermediate steps at a speed of 2 knots. The duration of tow varied from 6 to 25 minutes depending on depth and water clarity. The nets were thoroughly washed with a salt water hose and sampled organisms immediately preserved in 4% (buffered) formaldehyde in seawater. Data concerning regional and seasonal differences in copepod abundances within the Gulf will be published elsewhere. For an account of general plankton abundance in the Gulf, see ROTHLISBERG & JACKSON (1982). In addition to the Gulf collection, copepod samples have been analysed from collections taken in shelf waters of the Malay Peninsula during May-June 1984 (OTHMAN et al., 1987). That programme was part of a baseline study of a predeveloped oil-field lying between latitude 5o45' and 5°55'N and longitude 103o34 ' and 103°44'E in the South China Sea. Samples were taken from 20 stations using plankton nets having 0.64 m2 mouth area with mesh size of 142

COPEPODS OF THE GULF OF CARPENTARIA

am. The method of hauling was similar to that described above. We also added data from a surface tow in the Strait of Malacca, performed by the first author near Port Dickson on June 17th 1982, and data from samples taken by the second author in the Gladstone area (Northeast Australia) during July 1979. In all these programmes, adult copepods were identified to species level. Identifications of all species were based on comparisons with published descriptions. 3. RESULTS AND DISCUSSION 3.1. SYSTEMATIC SPECIES LIST A total of 102 species of copepods were recorded during the study, these including 68 species of calanoids from 34 genera and 22 families, 30 species of cyclopoids from six genera and four families and four species of harpacticoids from four genera in four families. The systematic list of these Gulf species as presented below follows the phylogenetic scheme of SARS (1903) for the calanoids, SARS (1913-1918) for the families of cyclopoids (with the genus Corycaeus after DAHL, 1912), and LANG (1948) for harpacticoids. It includes recommendations contained in revisions of the Calanidae by BRADFORD& JILLETT (1974), the Paracalanidae by ANDRONOV (1970, 1972a, 1972b), and the Candaciidae by GRICE (1963). In the Corycaeidae we include the species Corycaeus sewelli, first described from Moreton Bay in the dissertation by THWtN(1972) though not yet formally published. Parasitic or semiparasitic copepods (including cyclopoids belonging to the family Lichomolgidae), were not included in this study. ORDER COPEPODA Sub-Order Calanoida Family Calanidae Genus Calanus Leach, 1816 Calanus minor Claus, 1863 Genus Canthocalanus A. Scott, 1909 Canthocalanus pauper (Giesbrecht, 1888) Genus Cosmocalanus Bradford & Jillett, 1974 Cosmocalanus darwinii (Lubbock, 1860) Genus Undinula A. Scott, 1909 Undinula vulgaris (Dana, 1849) Family Eucalanidae Genus Eucalanus Dana, 1852 Eucalanus attenuatus (Dana, 1849) Eucalanus crassus Giesbrecht, 1888 Eucalanus dentatus A. Scott, 1909 Eucalanus mucronatus Giesbrecht, 1888 Eucalanus subcrassus Giesbrecht, 1888 Family Paracalanidae Genus Acrocalanus Giesbrecht, 1888 Acrocalanus gracilis Giesbrecht, 1888

563

Acrocalanus gibber Giesbrecht, 1888 Acrocalanus Iongicornis Giesbrecht, 1888 Genus Bestiola Andronov, 1972 Bestiola similis (Sewell, 1914) Genus Paracalanus Boeck, 1864 Paracalanus aculeatus Giesbrecht, 1888 Paracalanus denudatus Sewell, 1929 Genus Parvocalanus Andronov, 1970 Parvocalanus crassirostris (Dahl, 1894) Parvocalanus elegans Andronov, 1972 Parvocalanus latus Andronov, 1972 Family Calocalanidae Genus Calocalanus Giesbrecht, 1888 Calocalanus pavo (Dana, 1849) Calocalanus plumulosus (Claus, 1863) Family Clausocalanidae Genus Clausocalanus Giesbrecht, 1888 Clausocalanus furcatus (Brady, 1883) Clausocalanus minor Sewell, 1929 Family Aetideidae Genus Bradyidius Giesbrecht, 1897 *Bradyidius styliformis Othman & Greenwood, 1987 Family Euchaetidae Genus Paraeuchaeta A. Scott, 1909 Paraeuchaeta concinna (Dana, 1849) Family Phaennidae Genus Brachycalanus Farran, 1905 *Brachycalanus rothlisbergi Othman & Greenwood, 1988 Family Scolecithricidae Genus Scolecithricella Sars, 1903 Scolecithricella orientalis Mori, 1942 Family Diaixidae Diaixid gen. nov. *Diaixid sp. A *Diaixid sp. B 13iaixid sp. C Family Stephidae Genus Stephos T. Scott, 1892 *Stephos sp. nov. Family Centropagidae Genus Centropages Kr6yer, 1848 Centropages furcatus (Dana, 1849) Centropages gracilis (Dana, 1849) Centropages orsinfi Giesbrecht, 1889 Family Diaptomidae Genus Pseudodiaptomus Herrick, 1884 Pseudodiaptomus mertoni FrLichtl, 1923 Family Temoridae Genus Temora Baird, 1850 Temora discaudata Giesbrecht, 1892 Temora turbinata (Dana, 1849) Family Lucicutiidae Genus Lucicutia Giesbrecht, 1898 *Lucicutia sp. nov. Family Arietellidae

564

B.H.R. OTHMAN, J.G. GREENWOOD & P.C. ROTHLISBERG

Genus Metacalanus Cleve, 1901 Metacalanus aurivillii Cleve, 1901 Genus Paramisophria T. Scott, 1897 *Paramisophria sp. nov. Family Pseudocyclopidae Genus Pseudocyclops Brady, 1872 *Pseudocyclops kulai Othman & Greenwood, 1989 *Pseudocyclops minya Othman & Greenwood, 1989 Family Ridgewayiidae Genus Ridgewayia Thompson & A. Scott, 1903 *Ridgewayia flemingeri Othman & Greenwood, 1988 Family Candaciidae Genus Candacia Dana, 1846 Candacia bradyi A. Scott, 1909 Candacia catula (Giesbrecht, 1889) Candacia discaudata A. Scott, 1909 Genus Paracandacia Grice, 1963 Paracandacia truncata (Dana, 1849) Family Pontellidae Genus Calanopia Dana, 1852 Calanopia aurivillii Cleve, 1901 Calanopia australica Bayly & Greenwood, 1966 Calanopia elliptica (Dana, 1849) Genus Labidocera Lubbock, 1853 Labidocera acuta (Dana, 1849) Labidocera bengalensis Krishnaswamy, 1953 *Labidocera carpentariensis Othman, Greenwood & Fleminger, 1982 Labidocera dakini Greenwood, 1978 *Labidocera farrani Greenwood & Othman, 1979 Labidocera laevidentata (Brady, 1883) Labidocera minuta Giesbrecht, 1892 Labidocera kr6yeri (Brady, 1883) Genus Pontella Dana, 1846 Pontella securifer Brady, 1883 Pontella danae Giesbrecht, 1889 Genus Pontellina Dana, 1852 Pontellina morii Fleminger & Hulsemann, 1974 Genus Pontellopsis Brady, 1883 Pontellopsis scotti Sewell, 1932 Pontellopsis herdmani Thompson & A. Scott, 1903 Pontellopsis tasmanensis Greenwood, 1978 Family Acartiidae Genus Acartia Dana, 1846 Acartia amboinensis Carl, 1907 Acartia erythraea Giesbrecht, 1892 Acartia pacifica Steuer, 1915 Family Tortanidae Genus Tortanus Giesbrecht, 1898 Tortanus barbatus (Brady, 1883) Tortanus gracflis (Brady, 1883) Sub-Order Cyclopoida Family Oithonidae

Genus Oithona Baird, 1843 Oithona attenuata Farran, 1913 Oithona attenuata forma latithoracica Fr0chtl, 1923 Oithona rigida Giesbrecht, 1892 Oithona simplex Farran, 1913 Family Corycaeidae Genus Corycaeus Dana, 1845 Corycaeus (Corycaeus) speciosus Dana, 1848 Corycaeus (Ditrichocorycaeus) andrewsi Farran, 1911 Corycaeus (Ditrichocorycaeus) asiaticus F. Dahl, 1894 Corycaeus (Ditrichocorycaeus) dahfi Tanaka, 1960 Corycaeus (Ditrichocorycaeus) erythraeus Cleve, 1901 Corycaeus (Ditrichocorycaeus) lubbocki Giesbrecht 1982 Corycaeus (Ditrichocorycaeus) sewelli Thwin 1972 Corycaeus (Ditrichocorycaeus) subtilis M. Dahl, 1912 Corycaeus (Onchocorycaeus) agilis Dana, 1848 Corycaeus (Onchocorycaeus) catus F. Dahl, 1894 Corycaeus (Onchocorycaeus) pacificus F. Dahl, 1894 Genus Farranula Wilson, 1932 Farranula concinna (Dana, 1849) Farranula gibbula (Giesbrecht, 1891) Family Oncaeidae Genus Oncaea Philippi, 1843 Oncaea clevei FrL~chtl, 1923 Oncaea media Giesbrecht, 1892 Oncaea venusta Philippi, 1843 Family Sapphirinidae Genus Sapphirina Thompson, 1889 Sapphirina angusta Dana, 1849 Sapphirina gastrica Giesbrecht, 1891 Sapphirina ovatolanceolata-gemma Dana, 1849 Sapphirina nigromaculata Claus, 1863 Sapphirina opafina Dana, 1849 Sapphirina scarlata Giesbrecht, 1891 Sapphirina stellata Giesbrecht, 1891 Genus Copilia Dana, 1849 Copilia mirabilis Dana, 1849 Copilia quadrata Dana, 1849 Sub-Order Harpacticoida Family Ectinosomatidae Genus Microsetella Brady & Robertson, 1873 Microsetella rosea (Dana, 1848) Family Tachidiidae Genus Euterpina Norman, 1903 Euterpina acutifrons (Dana, 1852) Family Miraciidae Genus Macrosetella A. Scott, 1901

COPEPODS OF THE GULF OF CARPENTARIA

Macrosetella gracilis (Dana, 1848) Family Pseudopeltidiidae Genus Clytemnestra Dana, 1848 Clytemnestra scutellata Dana, 1848 The Gulf samples contained 13 species (marked with an asterisk (*)) which were new to science, of which descriptions have now been published for the following seven: Labidocera farrani GREENWOOD & OTHMAN, 1979, Labidocera carpentariensis (FLEMINGER et al., 1982), Bradyidius styliformis (OTHMAN & GREENWOOD, 1987), Brachycalanus rothlisbergi (OTRMAN & GREENWOOD, 1988a), Ridgewayia flemingeri (OTRMAN & GREENWOOD, 1988b), Pseudocyclops kulai (OTRMAN & GREENWOOD, 1989) and Pseudocyclops minya (OTRMAN & GREENWOOD 1989); the remaining six species we have denoted here without specific names, awaiting publication of our descriptions and taxonomic proposals. A further 10 species are recorded here for the first time from Australian waters, these are: Eucalanus dentatus,

Parvocalanus elegans, R latus, Scolecithricella orientalis, Labidocera bengalensis, Pontellopsis scotti, Acartia amboinensis, A. erythraea, Oithona simplex, and Oithona attenuata forma latithoracica. 3.2. PREVIOUS COPEPOD RECORDS FOR THE

GULF In his studies of zooplankton from waters to the north of Australia, which included examination of 93 samples from the Gulf, MARKINA (1972b) gave few details but noted that more than 100 copepod species were encountered. He identified only 20 of these species in his publication, hence no overall faunistic comparison can be made with his findings. It is noteworthy however that the only copepod species which MARKINA (1972b) specifically mentioned as occurring in the Gulf (Ioc. cit. pag. 9 '..particularly abundant in the ... outer parts of the Gulf..') was Acartia negligens, which was not found in our study. Of the remaining 19 species noted by Markina from seas north of Australia, only seven are amongst the species recorded from the Gulf in our study; two of these (Centropages furcatus, Labidocera acuta) were amongst the eight he characterised as common neritic forms around northern Australia. The remaining five (Undinula vulgaris, Undinula (= Cosmocalanus) darwinii, Euchaeta concinna, Eucalanus crassus and E. subcrassus) were amongst 14 referred to as tropical oceanic. Clearly A. negligens and many of the other species (identified and unidentified) encountered by Markina in waters near the north coast of Australia, comprise a list of species likely to be encountered in Gulf waters though not recorded in our studies. The 12 remaining species MARKINA (1972b) mentioned are:

Calanus lighti, Eucalanus subtenuis, Euchaeta con-

565

sirnilis, E. wolfendeni, E. marina, Centropages bradyi, C. gracilis, Lucicutia flavicornis, Labidocera detruncata, L. pavo, Pontellina plumata, and Acartia danae. 3.3. COMPARISON WITH THE COPEPOD FAUNA OF SOUTHEAST ASIA More detailed studies of copepod faunistics have been reported from eastern Indonesia by SCOTT (1909), FROCHTL (1923, 1924) and VERVOORT (1946). TO provide a basis for comparison with our findings, we have extracted species data from selected relevant station samples in these earlier studies. We selected 18 samples out of the 83 net catches studied by SCOTT (1909) from the Siboga Expedition of 1899-1900; these selected samples all came from stations east of 127°20'E (Buru-Ambon islands, Fig. 2), there being nine samples from the Seram Sea in August/September 1899 (station numbers 165 to 186) and nine from the Banda Sea in November 1899 to January 1900 (stn nos 225 to 282) (see Fig. 2 for localities). Scott recorded 194 copepod species from these 18 stations. FRUCHTL (1923) studied four samples collected by Merton in February-April 1908 around Aru, a group of islands at the transition of the deep Banda Sea and the shallow Arafura Sea, from which he recorded 72 copepod species. Finally, the copepods collected during the Snellius Expedition 1929-1930 were examined by VERVOORT(1946). In his study, only the results for the genera from Calanus up to and including Monacilla were given. For comparison with the present study, we selected Snellius stations from approximately those same areas as with Scott's study: i.e. the eastern Banda Sea plus the Seram Sea, but including the waters southeast of Timor and the Banda Arc where the Snellius performed one of her station legs. Twenty seven copepod species were recorded in 13 samples from October/November 1929 (stn nos ranging from 90 to 131) and three samples from April 1930 (stn nos 234 to 241) (VERVOORT, 1946). Pooling of the above lists shows that at least 72 of the 102 species reported here from the Gulf have also been found in the adjacent areas of eastern Indonesia (Table 1, left column). Twenty-five of the 30 remaining species have only been differentiated and described since the studies by SCOTT (1909) and FROCHTL (1923, 1924), and therefore could have occurred unrecognized in their collections; this is probable, since one, Labidocera carpentariensis, was collected near Aru during the Moro Expedition in July 1979 (FLEMINGERet al., 1982), and since another nine of those species were recently found further to the west along the coast of the Malay Peninsula (OTRMANet al., 1987), see Table 1. The five species all described before 1909 and taken in the Gulf but

566

B.H.R. OTHMAN, J.G. G R E E N W O O D & P.C. ROTHLISBERG

TABLE 1 Copepods of the Gulf of Carpentaria, their distribution in the adjacent and Australian waters and the major oceans (A = eastern Banda Sea area; B = coast Malay Peninsula; C = Great Barrier Reef; D = Moreton Bay area; E = off New South Wales coast; F = southern Australian coast; I = Indian Ocean; P = Pacific Ocean; At = Atlantic Ocean). Sources of data for the Southeast Asian and Australian areas: A = SCOTT (1909); FROCHTL (1923, 1924); VERVOORT(1946); plus 1 in sample Aru station (FLEMINGER et al., 1982). B = OTHMAN et al. (1987), plus 2 surface tow Port Dickson plus 4 sample from the Dulang area (Othman, unpublished data) and 3 NISHIDA (1985). C = FARRAN (1936, 1949); HODGSON (1982); WILLIAMS et al. (1988). D = THWIN (1972); GREENWOOD (1976, 1977, 1978a, 1978b, 1979). E = DAKIN & COLEFAX (1933, 1940); KOTT (1957); TAFE & GRIFFITHS (1983). F = BRADY (1883); NICHOLLS (1944); KOTT (1955); ARNOTT (1974a, 1974b). Sources of data for major oceanic areas: I = SEWELL (1914, 1929-1932, 1947); KRISHNASWAMY (1953); TANAKA (1960); ANORONOV (1972a); SILAS & PILLAI (1973); NISHIDA (1985). P = see C, D and E, plus MORI (1937, 1942); TANAKA (1956, 1957, 1964, 1965); GRICE (1962); MOTODA (1963); CHEN & ZHANG (1965); FROST & FLEMINGER (1968); FLEMINGER & HULSEMANN (1974); NISHIDA et al. (1977). 5 in sample from the Gladstone area, S.E. Queensland (Greenwood, unpublished data). At= GIESBRECRT (1892); T. SCOTT (1894); WILSON (1932); VERVOORT (1963, 1965); CERVIGON (1964); GONZALEZ & BOWMAN (1965); OWRE & FOYO (1967); PARK (1970). M a j o r Oceanic areas

Indo-Malay/Australian areas Species 1. 2. 3. 4. 5.

6. 7. 8. 9. 10. 11. 12. 13.

14. 15. 16.

17. 18. 19.

20. 21. 22. 23. 24. 25. 26. 27. 28. 29. 30. 31. 32. 33. 34. 35. 36. 37. 38. 39. 40.

Calanus m i n o r Canthocalanus pauper C o s m o c a l a n u s darwinfi U n d i n u l a vulgaris Eucalanus attenuatus E u c a l a n u s crassus Eucalanus dentatus Eucalanus mucronatus Eucalanus subcrassus A c r o c a l a n u s gracilis Acrocalanus gibber A c r o c a l a n u s Iongicornis Bestiola similis Paracalanus aculeatus Paracalanus denudatus P a r v o c a l a n u s crassirostris Parvocalanus elegans P a r v o c a l a n u s latus Calocalanus pavo Calocalanus plumulosus C l a u s o c a l a n u s furcatus Clausocalanus minor B r a d y i d i u s styliformis Paraeuchaeta concinna B r a c h y c a l a n u s rothlisbergi Scolecithricella orientalis

A

B

C

D

E

F

I

P

At

+ + + + + + + + + + + + + +

+ +

+ + + + + +

+

+

+

+

+

+

+ +

+ +

+ +

4+

+ +

+ +

+ + +

+ +

+

44+

+ + +

+ + +

+ + + + + + + + + + + + + + +

+ + + + + + + + +

+

+ + + +

+ + +

4-

4-

+ + + + + + + + +

+ + + + + + + +

+ + + +

+ + + + + + + + +

+

+

+

+

44-

+ + + +

4-

+

+

+ + +

4-

Diaixid sp. A Diaixid sp. B Diaixid sp. C S t e p h o s sp. nov.

C e n t r o p a g e s furcatus C e n t r o p a g e s gracilis C e n t r o p a g e s orsinfi Pseudodiaptomus mertoni Temora d i s c a u d a t a Temora turbinata Lucicutia sp. nov. M e t a c a l a n u s aurivillii P a r a m i s o p h r i a sp. nov. P s e u d o c y c l o p s kulai 41. P s e u d o c y c l o p s m i n y a 42. R i d g e w a y i a f l e m i n g e r i 43. C a n d a c i a b r a d y i

+ + +

+ + + + + +

+ + +

+ + + + +

+

+ + + +

4-

4-

+ + + + + +

+ + + + +

+

+ + + + + +

+5 + + + + + +

+

+

-I-

4-

+

+

COPEPODS

OF

THE

GULF

OF

CARPENTARIA

567

Indo-Malay/Australian areas Species 44. 45. 46. 47. 48. 49. 50. 51. 52. 53. 54. 55. 56. 57. 58. 59. 60. 61. 62. 63. 64. 65. 66. 67. 68. 69. 70. 71. 72. 73. 74. 75. 76. 77. 78. 79. 80. 81. 82. 83. 84. 85. 86. 87. 88. 89. 90. 91. 92. 93. 94. 95. 96. 97. 98. 99. 100. 101. 102.

Candacia catula Candacia discaudata Paracandacia truncata Calanopia aurivillii Calanopia australica Calanopia elliptica Labidocera acuta Labidocera bengalensis Labidocera carpentariensis Labidocera dakini Labidocera farrani Labidocera laevidentata Labidocera minuta Labidocera kro'yeri Pontella securifer Pontella danae Pontellina morii Pontellopsis scotti Pontellopsis herdmani Pontellopsis tasmanensis Acartia amboinensis Acartia erythraea Acartia pacifica Tortanus barbatus Tortanus gracilis Oithona attenuata O . att. f o r m a latithoracica Oithona plumifera Oithona rigida Oithona simplex Corycaeus (C.) speciosus Corycaeus (D.) andrewsi Corycaeus (D.) asiaticus Corycaeus (D.) dahfi Corycaeus (D.) erythraeus Corycaeus (D.) lubbocki Corycaeus (D.) sewelli Corycaeus (D.) subtilis Corycaeus (0.) agilis Corycaeus (0.) catus Corycaeus (0.) pacificus Farranula concinna Farranula gibbula Oncaea clevei Oncaea media Oncaea venusta Sapphirina angusta Sapphirina gastrica Sapphirina nigromaculata Sapphirina opafina Sapphirina ovatolanceolata Sapphirina scarlata Sapphirina stellata Copilia mirabilis Copilia quadrata Microsetella rosea Euterpina acutifrons Macrosetella gracilis Clytemnestra scutellata

A

B

C

D

+ + + +

-I+ -F + + + + +

-F + + +

-I+ + + + -F -t-

+ +

-I-t-

Major Oceanic areas E

F

+ -I-

-t-

I

P

+

+

+

+

+ +

+ +

+

+

+

+

+ +

+ +5

At

+1 +

+ + + +

+

-I-

-t-

,-J-

-I-

-F

+

+

-.F +2 -I+2 +2

-t+ -t-

-t-

+ -I-I+

-'F

-t-

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568

B.H.R. OTHMAN, J.G. GREENWOOD & P.C. ROTHLISBERG

not recorded in the collections of Scott and Fr0chtl, are Pontella danae (but recorded at a Siboga station on the north coast of Sulawesi), Pontellopsis herd-

mani, Acartia amboinensis, Corycaeus erythraeus and Clytemnestra scutellata. All these species have also recently been recorded along the Malaysian coast (column B, Table 1) and therefore presumably occur in intervening waters. They were not reported by SCOTT (1909), but his descriptions are so brief it is possible that some of these species did occur in the Siboga samples but were not differentiated. When the older species lists considered above are combined with the species list from OTHMAN et al. (1987) and some additional data (see Table 1), at least 88 out of the 102 species recorded in the Gulf are known to occur in Southeast Asian Waters, indicating a high level of faunal affinity in neritic copepods from the two regions. For the remaining 14 species missing in the records from Southeast Asian Waters, 11 belong to the new species discovered in the Gulf whereas the other three concern recently described species for the Pacific (Scolecithricella orientalis, 1942; Labidocera dakini, 1978; Pontellopsis tasmanensis, 1978). It is possible that all these species, though not found in the Malay Peninsula samples, do occur in eastern Indonesia, at least in the Arafura Sea. This faunal similarity is not surprising in view of the dynamics of water exchanges between the Gulf and surrounding water bodies. Early studies by ROCHFORD (1966) and NEWELL (1973) both concluded that large amounts of Coral Sea water entered the Gulf through the Torres Strait. Subsequently however, and based on improved data, FORBES (1984) calculated the exchange rates necessary to satisfy these conditions. He concluded that most seasonal changes in Gulf water occur in situ due to solar radiation, precipitation, wind and tidal mixing, and that these exchanges may be as important as exchanges with Arafura or Banda Sea water. Therefore, given the negligible mean flows calculated through the very shallow Torres Strait, most water exchange in the Gulf would have to come via the Arafura Sea through a much larger opening than the Torres Strait; namely through the Indonesian seas (WYRTKI, 1961; GODFREY & GOLDING, 1981 ; GODFREY & RIDGWAY, 1985; GORDON, 1986). The large number of species (about 100) previously recorded from eastern Indonesian waters, but not from the Gulf, are mainly meso- and bathypelagic forms which avoid shelf waters. Epipelagic species which spend part of their life cycle at greater depths were also absent from the Gulf samples. A striking example was the absence of Calanoides carinatus from the Gulf. This species was reported by SCOTT (1909), FROCHTL (1923, 1924) and VERVOORT (1946) from the eastern Banda Sea and adjacent waters

and is typical of upwelling areas, as shown by TRANTER (1977) for the waters south of Java. Presumably, all these records may concern a separate species C. philippinensis (KITOU& TANAKA, 1969, see also BRADFORD & JILLETT, 1974; FLEMINGER, 1985) as all the Calanoides specimens caught in eastern Indonesian waters during the Moro Expedition in 1979 belonged to this Indo-Pacific species (FLEMINGER,1986). During the southeast monsoon, upwelling occurs in the eastern Banda Sea (WYRTKI, 1961; ZIJLSTRA et al., 1990), and nutrientrich slope water has been reported to spread into the Arafura Sea close to the entrance of the Gulf by ROCHFORD (1966). However, subsequent studies (NEWELL, 1973; FORBES,1984) have failed to detect any penetration of this type of water in the Gulf and, similarly, no clear monsoonal influences on plankton biomass in the Gulf were found in the years 1975-1977 by ROTHLISBERG & JACKSON (1982). The effect of any such slope upwelling seems to be limited to the northern Arafura Sea and the Aru Basin, as indicated by much higher phytoplankton and zooplankton densities there during the southeast monsoon compared with the northwest monsoon (GIESKESet al., 1988; BAARS et al., 1990). In consequence C. philippinensis, which probably remains in diapause at depths of 300 m or more during the northwest monsoon, has only been found in the Aru Basin and other deep waters of eastern Indonesia, not in the shallow Arafura Sea proper (FLEMINGER, 1986; and recent collections during the Snellius-II Expedition, pers. comm. M.A. Baars). 3.4. COMPARISON WITH AUSTRALIAN RECORDS Previous studies of copepods in Australian waters have been concentrated along the east coast, especially in coastal waters of the Great Barrier Reef, southern Queensland (Moreton Bay), New South Wales, and to a lesser extent along the south Australian coast (Table 1). Of the 102 species found in the present study 63 species have been recorded from the Great Barrier Reef, 68 from Moreton Bay, 50 off New South Wales, and 17 off South Australia. Although FARRAN (1936) collected 193 copepod species in the Great Barrier Reef area, many of those species are regarded as open sea and deep water forms. Farran (l.c.) categorised the copepods into three groups of which 47 species were coastal, 100 species open sea, and 46 species were deep water. Considering only the coastal species, 2/3rds also occur in the Gulf. The greater number of species recorded in Moreton Bay, and shared with the Gulf, reflects the neritic nature of these areas, and that the sampling programmes carried out in Moreton Bay were more intensive than those in the Great Barrier Reef. If we combine the species lists for the Great

COPEPODS OF THE GULF OF CARPENTARIA

Barrier Reef and Moreton Bay, a total of 79 species from those areas are among the 102 reported here for the Gulf. Further south the number of species in common with the Gulf area decreases. The mechanism for dispersal into southern Queensland waters is the East Australian Current (EAC). North of Sydney (New South Wales) the EAC veers away from the coast and meanders east across the Tasman Sea (NILSSON & CRESSWELL, 1981). However, species of tropical origin have been found as far south as Tasmania, embedded in warmcore eddies that have been shed from the EAC. Of the 20 most abundant copepod species TRANTER et al. (1983) found in eddy J. (September 1979 - May 1980) in southern New South Wales, five (Oncea venusta, Calanus minor, Eucalanus attenuatus, Cosmocalanus darwinfi and Eucalanus crassus) were found in the Gulf. 3.5. RELATION TO INDIAN AND PACIFIC OCEANS The copepods sampled from the Gulf are mainly tropical/subtropical neritic species, however 40 are cosmopolitan and 17 have their distributional boundaries spread into more temperate areas. The Gulf fauna is approximately equally represented in both the Pacific and the Indian Oceans, and to a lesser extent in the Atlantic Ocean. Forty species have been recorded from the Atlantic, 86 from the Pacific, and 84 from the Indian Oceans. Three species recorded here from the Gulf (Parvocalanus elegans, Parvocalanus latus, Pontellopsis scottO have been previously recorded from the Indian Ocean but not from the Pacific. Conversely, five species taken in this study have been recorded from the Pacific Ocean, but not from the Indian Ocean, these are: Scolecithricella orientalis, Stephos sp. nov. (this same new species was recently collected by one of us in the Gladstone area, cf. Table 1), Labidocera dakini, Labidocera farrani and Pontellopsis tasmanensis. They all are neritic and have been described since the studies by SCOTT (1909) and FROCHTL (1923, 1924). The extent they may occur in Southeast Asian Waters is unknown, but, as we have said before, it is possible they are limited to the eastern part of the region, as none were recorded along the coast of the Malay Peninsula by OTHMAN et al. (1987). The two Parvocalanus species noted above, together with P. crassirostris, were very widespread in the Gulf. Parvocalanus crassirostris ranked third in overall abundance while the other two parvocalanids ranked seventh and eighth, and their ubiquity ranged from 40 to 70% (OTHMAN,1986). All three of these parvocalanids were subsequently found to be equally widespread and abundant in Malayan coastal waters (Othman, unpublished).

569

These three parvocalanids also share a common characteristic in being very small, with a range in body length of 0.35 to 0.55 mm. The paucity of records of P. crassirostris in Southeast Asia and western Pacific areas is very conspicuous and may be largely attributed to their minute size and hence possible escape through the relatively coarse nets commonly used in plankton studies, as suggested by GREENWOOD (1976). Further investigation is needed to establish whether both P. elegans and P. latus do occur in the western Pacific region. Biogeographically, it seems justifiable to regard the Gulf as belonging to the Indo-Pacific marine domain, forming the southeasternmost region. One of the species found in the Gulf is endemic to this region: Eucalanus dentatus. However this species, assumed to be neritic, has been caught, just south of Java 11°S 110°E; FLEMINGER, 1973; TRANTER, 1977), so could formally also be regarded to occur in the Indian Ocean). One of the 13 new species first discovered in the Gulf, Labidocera carpentariensis is most probably endemic also. After being recognized in the 1975 samples, the species subsequently has been found only in areas to the west of Cape York, as far as Darwin and Aru (FLEMINGERet al., 1982). Lucicutia sp. nov., the description of which is in press, has been recently found along the Malay Peninsula (Table 1) and is perhaps an endemic as well. In fact, the majority of the 13 new species from the Gulf might all have restricted distributions, as only two of them have been so far encountered in the Pacific (Northeast Australia, cf. Table 1), and none in the Indian Ocean. From the studies by SEWELL (1948) and FLEMINGER (1986) it is clear that Southeast Asia is an area with enhanced speciation in copepods, due to its geological past. Sea levels were lowered during the Pleistocene glacials and both the Sunda and Sahul shelves were exposed. The Banda Sea area then formed the only connection between the Indian and Pacific Oceans. FLEMINGER (1986) hypothesized that in this area upwelling was then much more pronounced than at present, hence stenothermal tropical copepods were isolated and evolved on either side of the cold water barrier. Opportunities for exchange of tropical copepod species between the two oceans was restored as sea levels rose after the last glacial, when other connections (including Torres Strait) were reestablished and the scale of upwelling diminished to become a brief seasonal phenomenon occurring only in a limited area. The occurrence in the Gulf of the five species previously only known from neritic waters of northeastern Australia may be an artefact of insufficient sampling, or reflect a geologically recent post-glacial invasion through Torres Strait. The relatively limited exchange rates possible through Torres Strait may

570

B.H.R. OTHMAN, J.G. GREENWOOD & P.C. ROTHLISBERG

explain why more species do not occur on both sides of Cape York. As a better understanding of PacificIndian Ocean water exchanges through the Indonesian Archipelago is developed, and more biogeographic resolution is added, a greater understanding of copepod species ranges and assemblages in this complex region will be gained. 4. REFERENCES

ALLDREDGE, A.L. & J.M. KING, 1977. Distribution, abundance, and substrate preferences of demersal reef zooplankton at Lizard Island Lagoon, Great Barrier Reef.--Mar. Biol. 41: 317-333. ANDRONOV, V.N., 1970. Some problems concerning the systematics of the family Paracalanidae (Copepoda).--Zool. Zh. 49: 980-985. (In Russian) ----, 1972a. Some new species of the genus Parvocalanus (Copepoda, Paracalanidae).--Zool. Zh. 51: 139-141. (In Russian) , 1972b. Veslonogie rachki Bestiola gen. n. (Copepoda, Paracalanidae).--Zool. Zh. 51: 290-292. (In Russian) ARNOTT, G.H., 1974a. Studies on the zooplankton of Port Phillip Bay and adjacent waters with special reference to Copepoda. Ph.D. Thesis, Monash University, Melbourne, Australia. , 1974b. An ecological study of the zooplankton of Westernport and adjacent Bass Strait Waters.-Ministry of Conservation, Vict., Aust., Publ. 95: 1-28. BAARS, M.A., A.B. SUTOMO, S.S. OOSTERHUIS & O.H. ARINARDI, 1990. Zooplankton abundance in the eastern Banda Sea and northern Arafura Sea during and after the upwelling season, August 1984 and February 1985.--Proc. Snellius-II Symp., Neth. J. Sea Res. 28: 527-543. BRADFORD,J.M. & J.B. JILLETT, 1974. A revision of generic definitions in the Calanidae (Copepoda, Calanoida).--Crustaceana 27: 5-16. BRADY, G.S., 1883. Report on the Copepoda collected by H.M.S. Challenger during the years 1873-76.Challenger Reports, Zool. 8: 1-142. CERVIGON, F., 1964. Los Corycaeidae del Caribe suroriental (Copepoda, Cyclopoida).--Mems Soc. Cienc. nat. La Salle 24: 163-201. CHEN, Q.-C. & S.-Z. ZHANG, 1965. The planktonic copepods of the Yellow Sea and the East China Sea. I. Calanoida.--Studia Mar. Sin. 7: 20-131. DAHL, M., 1912. Die Copepoden der Plankton-Expedition. I. Die Corycaeinen.--Plankton-Expedition der Humbold-Stiftung 2: 1-135. DAKIN,W.J. & A.N. COLEFAX,1933. The marine plankton of the coastal waters of New South Wales. I. The chief coastal forms and their seasonal distribution.--Proc. Linn. Soc. N.S.W. 58: 186-212. ----, 1940. The plankton of the Australian coastal waters off New South Wales. Part 1. Pubis Univ. Sydney Dep. Zool.: 1-211. FARRAN,G.P,, 1936. Copepoda.--Sci. Rep. Gt Barrier Reef Exped. 5: 73-142. ----, 1949. The seasonal and vertical distribution of the Copepods.--Sci. Rep. Gt Barrier Reef Exped. 2: 291-312.

FLEMINGER,A., 1973. Pattern, number, variability, and taxonomic significance of integumental organs (sensilla and glandular pores) in the genus Eucalanus (Copepoda, Calanoida).--Fishery Bull. 71: 965-1010. ,1985. Dimorphism and possible sex change in copepods of the family Calanidae.--Mar. Biol. 88" 273-294. ,1986. The Pleistocene equatorial barrier between the Indian and Pacific Oceans and a likely cause for Wallace's line.--Unesco Tech. Pap. Mar. Sci. 49: 84-97. FLEMINGER, A. & K. HULSEMANN, 1974. Systematics and distribution of the four sibling species comprising the genus Pontellina Dana (Copepoda, Calanoida).-Fishery Bull. 72: 63-120. FLEMINGER,A., B.H.R. OTHMAN& J.G. GREENWOOD, 1982. The Labidocera pectinata group: an Indo-West Pacific lineage of planktonic copepods with description of two new species.--J. Plankton Res. 4" 245-270. FORBES,A.M.G., 1984. The contribution of local processes to seasonal hydrology of the Gulf of Carpentaria.-Oceanogr. Tropicale 19: 193-201. FROST,B.W. & A. FLEMINGER,1968. A revision of the genus Clausocalanus (Copepoda: Calanoida) with remarks on distributional patterns in diagnostic characters.Bull. Scripps Instn Oceanogr. tech. Ser. 12: 1-235. FRUCHTL, F., 1923. Cladoceren und Copepoden der Aruinseln. Vorl&ufige Mitteilung: Artenliste und kurze Diagnosen der neuen Formen.--Abh. Senckenb. naturforsch. Ges. 35: 449-457. ----, 1924. Die Cladoceren und Copepoden - Fauna des Aru-Archipels.--Arb. zool. Inst. Univ. Innsbruck 2" 25-136. GIESBRECHT, W. 1892. Systematik und faunistic der pelagischen Copepoden des Golfes von Neapel. Fauna und Flora des Golfes von Neapel und der angrenzenden Meeresabschnitte: 1-831, 54 pls. GIESKES, W.W.C., G.W. KRAAY, A. NONTJI, O. SETIAPERMANA & SUTOMO, 1988. Monsoonal alternation of a mixed and a layered structure in the phytoplankton of the euphotic zone of the Banda Sea (Indonesia): a mathematial analysis of algal pigment fingerprints.-Neth. J. Sea Res. 22: 123-137. GODFREY,J.S. & T.J. GOLDING, 1981. The Sverdrup relation in the Indian Ocean, and the effect of PacificIndian Ocean throughflow on Indian Ocean circulation and on the East Australian Current.--& Phys. Oceanogr. 11 : 771-779. GODFREY,J.S. & K.R. RIDGWAY,1985. The large-scale environment of the poleward-flowing Leeuwin Current, Western Australia: Longshore steric height gradients, wind stresses and geostrophic flow.--J. Phys. Oceanogr. 15: 481-495. GONZALEZ, J.G. & T.E. BOWMAN, 1965. Planktonic copepods from Bahia Fosforescente, Puerto Rico, and adjacent waters.--Proc. U.S. natn Mus. 117: 241-303. GORDON, A.L., 1986. Interocean exchange of thermocline water.--J, geophys. Res. 91: 5037-5046. GREENWOOD, J.G., 1976. Calanoid copepods of Moreton Bay (Queensland) I. Families Calanidae to Paracalanidae.--Proc. R. Soc. Qd 87: 1-28. ----, 1977. Calanoid copepods of Moreton Bay (Queensland) I1. Families Calocalanidae to Centropagidae.-Proc. R. Soc. Qd 88: 49-67.

COPEPODS OF THE GULF OF CARPENTARIA

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