zootaxa

ZOOTAXA 62: 1-8 (2002) www.mapress.com/zootaxa/

ISSN 1175-5326 (print edition)

Copyright © 2002 Magnolia Press

ISSN 1175-5334 (online edition)

ZOOTAXA

New records and a checklist of continental free-living Copepoda (Crustacea) from Panama WILL K. REEVES1 & JANET W. REID2 1

Department of Entomology, Clemson University, Clemson, SC 29634, USA; email: [email protected]

2

Research Associate, Department of Science and Learning, Virginia Museum of Natural History, Martinsville, Virginia; Correspondence Address: 1100 Cherokee Court, Martinsville, VA 24112-5318, USA

Abstract We report new national records of Thermocyclops decipiens and Mesocyclops pseudomeridianus and two apparently undescribed species of Diacyclops and Halicyclops from Panama. We provide a complete bibliography of all literature on Panamanian copepods, and a table of historic copepod collection sites is presented. Key words: Copepoda, Panama, Crustacea, checklist

Introduction Microcrustaceans thrive in freshwater tropical habitats, where copepods are major components of planktonic assemblages (Covich & Thorp, 1991). The planktonic copepods, unlike the benthic or subterranean copepods, of Central America are reasonably well known. However, few collections have been made in freshwater habitats of Panama. There have been only two general copepod surveys in that country, by Marsh (1913) and Dodds (1926). These surveys were followed by subsequent reports on Panamanian material from the Marsh Collection in the National Museum of Natural History, Smithsonian Institution, U.S.A. (Coker, 1943; Weers & Zaret, 1975; Rocha, 1991; Reid, 1993; Reid & Mirabdullayev, 2000); and studies of plankton in the large artificial lakes of Gatún (Zaret, 1971, 1975, 1980, 1984; Zaret & Suffern, 1976; Garcés, 1982) and Bayano (Garcés, 1984). In the most recent regional checklist (Reid, 1990), only 17 of 57 species listed from Central America had been reported from Panama: 4 of 10 species of Calanoida, 12 of 35 Cyclopoida, and 1 of 12 Harpacticoida. Accepted: 26 July 2002; published: 8 August 2002

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Previous copepod collections were primarily from open waters of artificial lakes and ponds. Limited collections by W. Reeves from both planktonic and non-planktonic habitats resulted in four new Panamanian records, including two probable new species. Because no regional copepod checklist since that provided by Lindberg (1955) has included references to the primary literature, we provide this information in the checklist given in Table 1. Several of the identifications by Marsh (1913) and Dodds (1926) were of European or North American species that probably do not live in Central America; these records require verification. Re-examination of some of Marsh's material has already resulted in changes in identifications; moreover, some of the names used by Marsh (1913) and Dodds (1926) are now synonymized with other taxa. The questionable identifications and current synonymies are also indicated in Table 1.

Materials and Methods Collections were made in three different localities using a plankton net (mesh diameter 150 micrometers). The net was washed between collections to remove contaminants. All specimens were preserved in 95% ethanol. Voucher specimens of the copepods have been deposited in the United States National Museum of Natural History, Smithsonian Institution (USNM) and the Clemson University Arthropod Collection, U.S.A. (CUAC). Collecting methods, sites, and dates were as follows. On 18 June 1999 a plankton net was pulled behind a boat to sample the epilimnion of the inundated section of the Rio Majecito, Lake Bayano, Panama Province (9°4' N, 78°45' W). Samples were taken between 12:00 and 13:00 h at a depth of 0-1 meters. Flooding from rain had mixed the waters in the inlet the preceding day. The Rio Majecito flows into the Pacific Ocean. On 20 and 21 June 1999 samples of water and mud were collected from a permanent freshwater seep on Isla Grande, Colon Province (9°40' N, 79°35' W). Isla Grande is a volcanic island on the Caribbean Coast of Panama, with small first- and second-order streams, and is partially forested. The seep emerged from the volcanic bedrock approximately 9 meters above sea level. The seep had no plant cover, and was exposed to direct sunlight for at least half of the day. On 24 June 1999 a plankton net was used to sample a small tributary of the Colorado River, Chiriqui Province (8°46' N, 82°28' W). The tributary was approximately 100 centimeters wide and 10 centimeters deep where sampled. The bottom substrata included bedrock, gravel, and woody debris. The Colorado River drains into the Pacific Ocean.

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Results and discussion

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62 All species of copepods presently known from Panama are listed in Table I. Our collection provided four new country records. Thermocyclops decipiens (Kiefer) (Copepoda: Cyclopidae), although not previously recorded from Panama, is widespread in tropical and warm-temperate regions worldwide, especially in mesotrophic to eutrophic waterbodies (Reid, 1989). The record of Mesocyclops pseudomeridianus Defaye & Dussart (Copepoda: Cyclopidae) represents a significant range extension from the type locality in French Guiana (Defaye & Dussart, 1989). Diacyclops sp. and Halicyclops sp. (Copepoda: Cyclopidae) appear to be new taxa, and are presently under study by specialists.

Acknowledgements Fieldwork was partially funded by an E.W. King Memorial Grant from the Department of Entomology, Clemson University. We thank M. Dale, I. Dillon, R. Empson, D. Hawk, and M. Reeves for their assistance with fieldwork, M. Dillon and M. Dillon for providing logistical support and equipment in Panama, and P.H. Adler and J. Goforth for reviewing this manuscript. This is technical contribution 4644 of the South Carolina Agriculture and Forestry Research System.

References Coker, R.E. (1943) Mesocyclops edax (S. A. Forbes), M. leuckarti (Claus) and related species in America. Journal of the Elisha Mitchell Scientific Society, 59, 181-200. Covich, A.P. & Thorp, J.H. (1991) Crustacea: introduction and Peracarida. In: Thorp, J.H. & Covich, A.P. (eds.), Ecology and Classification of North American Freshwater Invertebrates. Academic Press, San Diego. pp. 593-664. Defaye, D., & Dussart, B.H. (1989). Compléments à la faune des crustacés copépodes des eaux intérieures de Guyane française. Revue d'Hydrobiologie Tropicale, 21, 109-125. Dodds, G.S. (1926) Entomostraca from the Panama Canal Zone. Occasional Papers of the Museum of Zoology of the University of Michigan, 8(174), 24-27. Garcés B., H.A. (1982) Composición y abundancia de los copépodos y rotíferos en el Lago Gatún. ConCiencia, 9, 12-15. Garcés B., H.A. (1984) Estudio preliminar sobre el zooplancton del Lago Bayano, Panamá. Revista de Biología Tropical, 32, 17-24. Lindberg, K. (1955) Cyclopoïdes (Crustacés copépodes) du Mexique. Arkiv för Zoologi, 7, 459489. Marsh, C.D. (1913) Report on fresh-water Copepoda from Panama. Smithsonian Miscellaneous Collections, 61, 12-19. Reid, J.W. (1989) The distribution of species of the genus Thermocyclops (Copepoda, Cyclopoida) in the western hemisphere, with description of T. parvus, new species. Hydrobiologia, 175, 149-174.

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Reid, J.W. (1990) Continental and coastal free-living Copepoda (Crustacea) of Mexico, Central America and the Caribbean region. In: Navarro L., D. & Robinson, J.R. (eds.), Diversidad Biologica en la Reserva de la Biosfera de Sian Ka'an, Quintana Roo, México. Centro de Investigaciones de Quintana Roo (CIQRO) and Program of Studies in Tropical Conservation, University of Florida; Chetumal, Quintana Roo. pp. 175-213. Reid, J.W. (1993) New records and redescriptions of American species of Mesocyclops and of Diacyclops bernardi (Petkovski, 1986) (Copepoda: Cyclopoida). Bijdragen tot de Dierkunde, 63, 173-191. Reid, J.W. & Mirabdullayev, I.M. (2000) First record from Panama of the freshwater copepod Yansacyclops ferrarii (Cyclopoida: Cyclopidae). Nauplius, 7, 187-189. Rocha, C.E.F. 1991. A new species of Halicyclops (Copepoda, Cyclopidae) from California, and a revision of some Halicyclops material in the collections of the US Museum of Natural History. Hydrobiologia, 226, 29-37. Walter, T.C. (1989) Review of the New World species of Pseudodiaptomus (Copepoda: Calanoida), with a key to the species. Bulletin of Marine Science, 45, 590-628. Weers, E.T., & Zaret, T.M. (1975) Grazing effects on nannoplankton in Gatún Lake, Panama. Verhandlungen der Internationale Vereinigung für Theoretische und Angewandte Limnologie, 19, 1480-1483. Zaret, T.M. (1971) Studies on Predator-prey Interactions in a Tropical Lacustrine Ecosystem. Ph.D. Thesis, Yale University, New Haven. 94 pp. Zaret, T.M. (1975) Strategies for existence of zooplankton prey in homogeneous environments. Verhandlungen der Internationale Vereinigung für Theoretische und Angewandte Limnologie, 19, 1484-1489. Zaret, T.M. (1980) Predation and Freshwater Communities. Yale University Press, New Haven. 187 pp. Zaret, T.M. (1984) Central American limnology and Gatún Lake, Panamá. In Taub, F.B. (ed.), Lakes and Reservoirs. Ecosystems of the World 23. Elsevier, Amsterdam. pp. 447-465. Zaret, T.M. & J.S. Suffern (1976) Vertical migration in zooplankton as a predator avoidance mechanism. Limnology and Oceanography, 21, 804-813.

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TABLE 1. Copepod crustaceans recorded from continental fresh and brackish water habitats in Panama. Species

Locality

Reference

Diaptomus s.l. gatunensis Marsh, 1913

Black Swamp near old Panama Railroad, and pond at Bohio, 40 samples including Gatún Lake, ponds, Miraflores Lake (brackish water), Lake Bayano

Marsh (1913), Dodds (1926), Zaret (1971, 1975, 1980, 1984), Weers & Zaret (1975), Zaret & Suffern (1976), Garcés (1982, 1984), present report (USNM 288062)

Diaptomus s.l. leoninicollinus Marsh, 1913

Lion Hill, Canal Zone, 64 samples including Gatún Lake, small ponds, Panama Canal

Marsh (1913), Dodds (1926)

Prionodiaptomus colombiensis (Thiébaud, 1912)

Comacho Reservoir and Rio Trinidad 7 samples from ponds between Gamboa and Summit

Marsh (1913, as Diaptomus marshi Juday, 1913), Dodds (1926, as D. marshi)

Pseudodiaptomus cristobalensis Marsh, 1913

Old French Canal

Marsh (1913), Walter (1989)

Pseudodiaptomus culebrensis Marsh, 1913

Rio Culebra, branch of Rio Chepo, 11 samples from Miraflores Lake (brackish water)

Marsh (1913), Walter (1989), Dodds (1926)

Pseudodiaptomus panamensis Walter, 1989

Naos Marine Station, Panama Canal (i.e., Pacific coastline)

Walter (1989)

Savannas between Panama and Old Panama City, 19 samples from Miraflores Lake (brackish water)

Marsh (1913, as Cyclops panamensis), Dodds (1926, as C. panamensis)

Order Calanoida

Order Cyclopoida Apocyclops panamensis (Marsh, 1913)

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TABLE 1 continued.

62 Species

Locality

Reference

Cyclops sp.6

Gatún Lake

Garcés (1982)

Diacyclops sp.4

Waterfall, Chiriquí, trib- Present report (USNM utary of the Colorado 288059) River

Ectocyclops phaleratus (Koch, 1838)1

Black Swamp, 12 sam- Marsh (1913, as ples from shore zones of Cyclops quinquepartitus Marsh, 1913), lakes and ponds Dodds (1926, as Cyclops phaleratus)

Eucyclops agilis (Koch, 1838)1

Comacho Reservoir, pond at Bohio, Black Swamp, Rio Trinidad, Miraflores, and savannas near Panama City, 39 samples from shores and bottoms of lakes and ponds, Gatún Lake

Marsh (1913, as Cyclops serrulatus Fischer, 1851), Dodds (1926, as C. serrulatus), Zaret (1971, 1975, 1980, 1984)

Halicyclops cf. clarkei Herbst, 19822

Savannas and a well near Panama City

Marsh (1913, as Cyclops aequoreus Fischer, 1860), Rocha (1991)

Halicyclops sp.4

Freshwater seepage on Isla Grande, Colón

Present report (USNM 288060)

Homocyclops ater (Herrick, 1882)

2 samples from pond near Darien and Miraflores Lake

Dodds (1926, as Cyclops ater)

Macrocyclops albidus (Jurine, 1820)

Black Swamp, 13 samples from weedy ponds and marshes

Marsh (1913, as Cyclops albidus) Dodds (1926, as C. albidus)

Order Cyclopoida


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TABLE 1. continued.

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62 Species

Locality

Reference

Mesocyclops leuckarti (Claus, 1857)1

Reservoirs and most samples, Canal Zone 85 samples from Agua Clara Reservoir, ponds and marshes, Lake Bayano

Marsh (1913, as Cyclops leuckarti) Dodds (1926, as C. leuckarti), Garcés (1984, as C. leuckarti)

Mesocyclops longisetus var. curvatus Dus-

Panama

Reid (1993) from Marsh Collection

Lake Bayano

Present report (USNM 288061)

Microcyclops anceps (Richard, 1897)1

Savannas between Panama and Old Panama City, 28 samples from ponds, marshes, and weedy shores of lakes, including near Frijoles

Marsh (1913, as Cyclops dentatimanus Marsh, 1913), Dodds (1926, as C. dentimanus)

Microcyclops varicans (G. O. Sars, 1863)1

Black Swamp

Marsh (1913, as Cyclops varicans)

Thermocyclops decipiens (Kiefer, 1929)4

Lake Bayano

Present report (CUAC)

Thermocyclops inversus (Kiefer, 1936)

Miraflores, pond near Bohio, savannas near Panama City, Gatún Lake; Rio Trinidad, Agua Clara, Calabraras, Mindi, Camacho, Cocoli, Carabali, and

Marsh (1913, as Cyclops tenuis in part), Coker (1943), Reid (1989)

Order Cyclopoida

3

sart, 1987

Mesocyclops pseudomeridianus Defaye & Dussart, 19894

Gorgona Reservoirs5 Thermocyclops tenuis (Marsh, 1909)

As for T. inversus5, 81 samples including Gatún Lake, ponds, Ft. Lorenzo reservoir

Marsh (1913, as C. tenuis in part), Coker (1943), Reid (1989), Dodds (1926, as C. tenuis)




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TABLE 1 continued.

62 Species

Locality

Reference

Tropocyclops prasinus (Fischer, 1860)1

Black Swamp, Gatún Lake, stream near Alhajuela caves, pond near Bohio, Miraflores Lake (brackish water)

Marsh (1913, as Cyclops prasinus) Dodds (1926, as C. prasinus)

Yansacyclops ferrarii Reid, 1988

Fresh water pond

Reid & Mirabdullayev (2000)

Canthocamptus staphylinus Jurine, 18201

Small stream near Gold Hill

Dodds (1926)

Canthocamptus spp.1

Small streams

Dodds (1926)

Order Cyclopoida

Order Harpacticoida

1

Doubtful record; to be taken as some species of that genus or family.

2

Re-identification of specimens reported as Cyclops aequoreus by Marsh (1913).

3

Re-identification of specimens reported as Cyclops leuckarti by Marsh (1913).

4

New record for Panama.

5

Some records may refer to other species of Thermocyclops (see Coker, 1943; Reid, 1989).

6

Cyclops is a boreal genus; this record should be taken to refer to some species of cyclopoid.

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