2008 SOUTHEASTERN NATURALIST 7(2):219–228 New Records of Copepods (Crustacea) from the Florida Keys Lawrence J. Hribar1,* and Janet W. Reid 2 Abstract - New records of copepods are listed from artificial containers and shallow waterbodies in the Florida Keys. Mesocyclops ogunnus is reported from the United States for the first time. Metacyclops cf. gracilis is reported from the Florida Keys for the first time. New collection records are presented for 12 other species. Mesocyclops ogunnus is an Old World species, and this is the third record in the Neotropics; it may have the potential to compete with the North and Central American native Mesocyclops edax. The non-native copepod species found in central and southern Florida are generally associated with plants and soils. A review of literature on introduced copepods found in South Florida is presented. Introduction The Florida Keys are islands that lie east, south, and southeast of peninsular Florida (Fig. 1). They are part of the South Florida rockland ecosystem, with a fl ora composed of both temperate and tropical components (Snyder et al. 1990, Stern and Brizicky 1957). We recently published a list of new records of copepods (Crustacea) from the Florida Keys, Monroe County, FL (Reid and Hribar 2006). Five species were reported from the United States for the first time, 6 species were reported from Florida for the first time, and new distribution records within Florida were provided for a further 19 species. Since the publication of those records, further collections have revealed additional distribution records for copepods in the Florida Keys, including one species reported from the United States for the first time, and another species previously known only from peninsular Florida. Materials and Methods The specimens reported herein were collected incidentally during routine surveillance for larval mosquitoes conducted by the Florida Keys Mosquito Control District. Samples of water containing mosquitoes were collected from potential habitats with the aid of 236-ml (half-pint) dippers or turkey basters (a large pipette fitted with a squeeze bulb), and returned to the laboratory (Service 1993). Copepods were removed with a pipette and preserved in 80% isopropanol and glycerin. Most collections were made by the first author; all specimens were identified by the second author. Taxonomic identification was made from specimens temporarily transferred to nearly full-strength glycerin (by gradual evaporation of a 70% isopropanol-10% glycerin mixture) with a small amount of 80% lactic acid added to the mixture. The specimens were examined by 1Florida Keys Mosquito Control District, 503 107th Street Gulf, Marathon, FL 33050. 2Virginia Museum of Natural History, 21 Starling Avenue, Martinsville, VA 24112. *Corresponding author - lhribar@keysmosquito.org. 220 Southeastern Naturalist Vol.7, No. 2 manipulation on a glass depression slide as described by Reid (2000), with the use of a Leica DMLB compound microscope, at magnifications of 400 or 800 x. The principal references used were the keys and descriptions by Bruno et al. (2005), Holyńska et al. (2003), and Wilson and Yeatman (1959). It proved unnecessary to dissect any specimens. The whole specimens were transferred to 70% isopropanol for long-term storage and were deposited in the Recent Invertebrates Collection of the Virginia Museum of Natural History (VMNH), Martinsville, VA. The collection numbers provided are from the VMNH Crustacea Catalog Database. Results: List of Species Order Calanoida, Family Diaptomidae Arctodiaptomus dorsalis (Marsh): 1F, Long Key, rain pool, 27 Feb 2007, coll. L. Hribar, VMNH 1437. This is a generalist Neotropical species with a core range from extreme northern South America through eastern Mexico and the Antilles, and into the southern USA; outlying records range from the central Colombia highlands and Venezuela to California, Arizona, the central United States as far north as Michigan, and the eastern United States as far as the District of Columbia, where the species is thought to have been transported along with stocked fish or ornamental aquatic plants (Reid 2008). It is widespread in peninsular Florida, especially in eutrophic ponds and lakes (Bruno et al. 2005), and was previously recorded from Grassy Key in the Florida Keys (Reid and Hribar 2006). New locality record. Order Cyclopoida, Family Cyclopidae Apocyclops dimorphus Kiefer: 4F, 2M, 55 copepodids, Lower Matecumbe Key, rain pool, 17 Apr 2007, coll. L. Hribar, VMNH 1438; 48F, 14M, 6 Figure 1. The Florida Keys, indicating the keys where samples were taken. 2008 L.J. Hribar and J.W. Reid 221 copepodids, Vaca Key, semi-permanent rain pond, 31 Oct 2007, coll. L. Hribar, VMNH 1481. This species is seldom collected but widely distributed in brackish to saline waters. It was originally described from Haiti, and is also known from the Texas coast and inland saline lakes in California and Mexico, and Big Pine Key and Vaca Key in the Florida Keys (Reid and Hribar 2006, Reid et al. 2002). New locality records. Apocyclops panamensis (Marsh): 50F, 50M, Vaca Key, boat, 11 Apr 2006, coll. E. Posada, VMNH 1439; 3F, 2M, 4 copepodids, Vaca Key, bird bath, 29 Jan 2007, coll. C. Samul, VMNH 1440; 15F, 2M, 13 copepodids, Fat Deer Key, rain pool, 20 Feb 2007, coll. L. Hribar, VMNH 1441; 4F, 1M, Fat Deer Key, rain pool, 17 Apr 2007, coll. L. Hribar, VMNH 1442; 1F, 3M, 3 copepodids, Fat Deer Key, mangrove swamp, 3 May 2007, coll. L. Hribar, VMNH 1443; 12F, 7M, 5 copepodids, Big Pine Key, buttonwood swamp, 30 May 2007, coll. L. Hribar, VMNH 1444. Apocyclops panamensis is common in brackish to hypersaline coastal habitats from the Atlantic coast of the United States to northern South America. Previous collections from the Florida Keys include Cudjoe Key, Key Largo, Long Key, Long Point Key, Stock Island, Sugarloaf Key, Vaca Key, and Windley Key (Reid and Hribar 2006, Yeatman 1963). The collections from Big Pine Key and Fat Deer Key are new locality records. Bryocyclops muscicola (Menzel), 2F, 1M, Key Largo, metal container, 2 Aug 2007, coll. J. Davis, VMNH 1445. This species was previously collected from a bromeliad on Duck Key (Reid and Hribar 2006). This is a new locality record, and apparently the first record of this species from an artificial container. Diacyclops bernardi (Petkovski), 1F, 1M, Big Pine Key, buttonwood swamp, 30 May 2007, coll. L. Hribar, VMNH 1446. Previous Florida Keys collection localities for this species are Key Largo, Long Point Key, and Windley Key (Reid and Hribar 2006). New locality record. Halicyclops sp. A: 1F, 6M, Grassy Key, mangrove swamp, 22 Feb 2007, coll. L. Hribar, VMNH 1447; 1F, Big Pine Key, buttonwood swamp, 30 May 2007, coll. L. Hribar, VMNH 1448. Members of this taxon have previously been collected from Key Largo (Reid and Hribar 2006). The present collections represent new locality records within the Florida Keys. The specimens do not match descriptions of any known Halicyclops species (Reid and Hribar 2006). Halicyclops sp. B: 1F, Long Key, rain pool, 1 Nov 2007, coll. L. Hribar, VMNH 1482. This morph, not collected before in the Florida Keys, does not correspond with previously described species from the Americas. Macrocyclops albidus (Jurine): 1M, 6 copepodids, Stock Island, swale ditch full of cattails (Typha sp.), 30 May 2007, coll. L. Hribar, VMNH 1449. This taxon is a common, eurytopic species, previously collected from Stock Island (Reid and Hribar 2006). Mesocyclops ogunnus Onabamiro: 3F, Vaca Key, plant tray, 4 Jun 2007, coll. C. Samul, VMNH 1450; 2F (one of which was bearing egg sacs), Vaca Key, semi-permanent rain pond, 31 Oct 2007, coll. L. Hribar, VMNH 1483. This species is normally planktonic, of Afro-Asian origin, and has been found in the New World in reservoirs in Brazil (Reid and Pinto-Coelho 1994a, b) and ponds in the Cayman Islands (Suárez-Morales et al. 1999). 222 Southeastern Naturalist Vol.7, No. 2 This record is the first report of M. ogunnus from the United States, and the first from an artificial container. Metacyclops cf. gracilis (Lilljeborg), 15F, 1M, Long Key, rain pool, 17 Apr 2007, coll. L. Hribar, VMNH 1451. Members of this taxon were collected previously in small numbers in the Florida Everglades (Bruno et al. 2005). It resembles the European M. gracilis except in minor morphological details, as discussed by Bruno et al. (2005). New Florida locality record. Microcyclops rubellus (Lilljeborg): 7F, 4M, 5 copepodids, Vaca Key, rain pool, 20 Feb 2007, coll. L. Hribar, VMNH 1452; 5F, 3 copepodids, Long Key, rain pool, 27 Feb 2007, coll. L. Hribar, VMNH 1453; 6F, 3M, Long Key, rain pool, 17 Apr 2007, coll. L. Hribar, VMNH 1454. This widely distributed species was previously known from Big Pine Key, Cudjoe Key, Duck Key, Long Key, Long Point Key, and Stock Island (Reid and Hribar 2006). The Vaca Key collection represents a new locality record within the Florida Keys. Order Harpacticoida, Family Ameiridae Nitokra lacustris (Shmankevich): 2F, 1M, Fat Deer Key, rain pool, 20 Feb 2007, coll. L. Hribar, VMNH 1455; 9F, 2M, 5 copepodids, Vaca Key, rain pool, 20 Feb 2007, coll. L. Hribar, VMNH 1456; 1F, Long Key, rain pool, 17 Apr 2007, coll. L. Hribar, VMNH 1457; 1F, Long Key, rain pool, 1 Nov 2007, coll. L. Hribar, VMNH 1485. This widely distributed species of coastal brackish to fresh waters previously was known from Key Largo, Long Point Key, and Stock Island. New locality records. Cletocamptus fourchensis Gómez, Fleeger, Rocha-Olivares, and Foltz: 5F, 5M, 3 copepodids, Vaca Key, semi-permanent rain pond, 31 Oct 2007, coll. L. Hribar, VMNH 1484. This species, known from coastal brackish waters in Louisiana and Mississippi, was reported from Vaca, Long, and Grassy keys by Reid and Hribar (2006). Family Ectinosomatidae Pseudectinosoma cf. minor (Kunz): 2F, Grassy Key, mangrove swamp, 22 Feb 2007, coll. L. Hribar, VMNH 1458. Members of this taxon were previously collected from Key Largo (Reid and Hribar 2006). As we discussed previously, the North American specimens are only provisionally assigned to this taxon; they must be compared to European material. The European P. minor, however, must be redescribed before any such comparisons can be made. New locality record. Discussion Southern Florida may be predisposed to invasion by exotics because of a number of factors, including insularity, relatively young geological age, numerous international ports of entry, subtropical climate, large-scale development, and an abundance of diverse, remote, and unmonitored freshwater habitats (Simberloff 1997, Warren 1997). Southern and central Florida indeed appears to be a “hot spot” for introduced copepods. Reid and Hribar (2006) reported 2 species, Bryocyclops muscicola (Menzel) and Paracyclops 2008 L.J. Hribar and J.W. Reid 223 bromeliacola Karaytug et Boxshall, which appear to have been introduced into Florida via human agency. Both of these species were detected in bromeliads. Bryocyclops muscicola was previously reported from organic soils of ornamental plants obtained by P.S. Lehman at two nurseries in Orange and Lake counties in central Florida (Bruno et al. 2005, Reid 1999). This tiny cyclopoid was originally described from Java and was later found in Sumatra; it is therefore presumed to be Asian in origin, although it most closely resembles two Brazilian species (Reid 1999). Paracyclops bromeliacola was described from leaf cups of arboreal and terrestrial bromeliads and leaf litter in Atlantic Forest sites in the state of São Paulo, Brazil (Karaytug and Boxshall 1998). Several hundred bromeliad species belonging to at least 48 genera have been imported into Florida (Cathcart 1995), and this importation of plants may be responsible for introduction of non-native species into the state (Grogan and Hribar 2006, Zavortink and O’Meara 1999). The collections reported here have added a third non-native cyclopoid, Mesocyclops ogunnus, in this case also associated with a plant container, as well as a semi-permanent rain pond. However, in contrast to B. muscicola and P. bromeliacola, M. ogunnus normally occurs in the plankton of freshwater to oligohaline lakes and reservoirs. Mesocyclops ogunnus is one of the most widely distributed Old World species of the genus; its range includes most of Africa and extends through Asia eastward as far as Japan (Kyushu, approximately 31°30'N; Ishida 2002), and northward to Uzbekistan (Holyńska et al. 2003) and Kazakhstan (approximately 45°N; Krupa 2005). Mesocyclops ogunnus is known from two other locations in the Americas, and is considered to be introduced in both. In central Brazil, it was first reported (as Mesocyclops kieferi van de Velde) in 1988, in Barra Bonita Reservoir in the state of São Paulo, where it had not previously been found as recently as 1985–1986, but in 1988 was present in abundance (Matsumura-Tundisi et al. 1990, Tundisi et al. 1991). It was then reported (as M. ogunnus) from two locations in Furnas Reservoir, on the border of São Paulo and Minas Gerais (Reid and Pinto-Coelho 1994a, b). In spite of the presence of several native congeners, M. ogunnus is expanding its range in reservoirs in central Brazil, in the cascade reservoirs of the Tietê /Upper Paraná river basin (Matsumura- Tundisi and Silva 2002). At last report, it was found only rarely in natural, lentic and lotic habitats of the Upper Paraná River fl oodplain, in the states of Paraná and Mato Grosso do Sul (Lansac-Tôha et al. 2002). In the eutrophic Barra Bonita Reservoir, it is now a codominant cyclopoid with the native Thermocyclops decipiens (Kiefer), as reported by Santos-Wisniewski and Rocha (2007). Mesocyclops ogunnus was also reported from the Cayman Islands, where it was found in two shallow coastal ponds in salinities of 2.6 and 0.8‰ (Suárez-Morales et al. 1999). Although Suárez-Morales et al. (1999) considered M. ogunnus to be an invader in the Caymans and noted that further sampling could better define its distribution on these islands, to our knowledge this follow-up sampling has not been done. Mesocyclops ogunnus is able to survive and reproduce at salinities up to 5‰, although it reaches a smaller body size in more-saline waters (Bonou et al. 1991). In the rain pond on Vaca Key, two females, one of which was carrying egg sacs, occurred together 224 Southeastern Naturalist Vol.7, No. 2 with Apocyclops dimorphus and Cletocamptus fourchensis; both of the latter species usually occur in brackish to hypersaline waters. We do not know the origin of the copepods found in the plant tray or the rain pond. In view of its euryhaline nature, M. ogunnus may easily be able to survive in the often brackish, shallow surface waters in the Keys, as evidenced by the ovigerous female present in the rain pond. Collections in likely habitats are continuing. In North America, we speculate that M. ogunnus may be able to successfully compete with the indigenous Mesocyclops edax (S.A. Forbes). The geographical distribution of M. edax extends from Nicaragua and Cuba, north through Mexico and the United States to central Canada (Reid and Moreno 1999), where it occurs up to approximately 61°N and in lakes with ice-free periods of about 135 days and mean July air temperatures over 15 °C, which allows water temperatures in the epilimnion to reach 20 °C in late July and early August (Patalas 1986). Both species live mainly in permanent lakes and ponds, and both have a predilection for meso- to eutrophic environments (e.g., Dobrzykowski and Wyngaard 1993, Santos-Wisniewski and Rocha 2007). In particular, M. ogunnus is highly successful in eutrophic impoundments; in parts of West Africa, it now is a dominant zooplankter in such environments (Pagano et al. 2003). Both species utilize the entire water column, although, like many planktonic copepods, they migrate diurnally, moving vertically upward at night and downward in daytime (see the review by Williamson 1986; for the pattern of M. ogunnus in Lake Kinneret, see Gophen 1978, who reported it as Mesocyclops leuckarti (Claus)). This migration is thought to be a means of escaping visually oriented predators, particularly fish. In Lake Kinneret, increased predation by Mirogrex (= Acanthobrama) terraesanctae (Goren) during 1972–75 may have depressed the numbers of adult M. ogunnus (even though cladocerans rather than copepods are preferred prey of this sardine), but fecundity (number of eggs per female) of the copepods increased fivefold in the same period (Gophen and Landau 1977). Bonou et al. (1991) examined aspects of the growth and development of individuals of M. ogunnus collected from fishponds in the Ivory Coast, finding that in laboratory cultures, development from egg to adult required a mean of 8.11 days at 30 °C; they noted that this rate was somewhat slower than rates measured for the same species in large freshwater lakes, and attributed the difference to nutritional conditions. In laboratory cultures, males of M. ogunnus from Lake Kinneret, Israel, matured from egg to adult in 52, 25, and 19 days at 15, 22, and 25 °C, respectively; females matured slightly more slowly (Gophen 1976, as M. leuckarti). In tropical and subtropical regions, M. edax persists in the water column year-round; but at least from Virginia northward, it passes the winter as diapausing older copepodids and adult females (viz. Dobrzykowski and Wyngaard 1993). Like most members of their genus, adults and older copepodids of both species are omnivorous selective predators that will take a wide range of food items, including cladocerans, copepods (especially nauplii and small copepodids), rotifers, and algae; see for example the reports of Confer (1971), Williamson (1980, 1984), and Janicki and DeCosta (1990) on M. edax, and of Gophen (1977) and Blumenshine and Hambright (2003) on M. ogunnus. The nauplii and younger copepodids are mainly herbivorous (Gophen 1977). 2008 L.J. Hribar and J.W. Reid 225 Different populations may show contrasting prey preferences; for example, Williamson (1980) found that M. edax avoided the cladoceran Bosmina longirostris (O.F. Müller), and Confer (1971) found that unstarved adults of M. edax preferred copepodids of Arctodiaptomus fl oridanus (Marsh) over cladocerans; whereas Janicki and DeCosta (1990) found that M. edax preferred B. longirostris to the cladoceran Daphnia parvula Fordyce and the calanoid Skistodiaptomus pallidus (Herrick). Under field conditions, M. ogunnus will predate aggressively upon larvae of the mosquito Aedes aegypti (L.); whereas M. edax will take larvae of several species of Aedes and Anopheles quadrimaculatus Say, but in small numbers (Marten and Reid 2007 and references therein). Havel et al. (2005) argued that the proliferation of reservoirs over the past century has contributed to the dispersal of native and exotic aquatic species because reservoirs are disturbed and variable habitats, are often eutrophic, and in arid areas are often more saline than natural lakes, and because their locations on rivers provide physical stepping stones between waterbodies, among other factors. The demonstrated affinity of M. ogunnus for artificial, eutrophic, and saline waterbodies may eventually allow this species to establish itself in the thousands of impoundments in the southeastern United States. Most water samples that were examined contained only one species of copepod. However, in addition to the sample from the semi-permanent rain pond on Vaca Key that contained M. ogunnus, A. dimorphus, and C. fourchensis, one sample taken from a rain pool on Long Key on April 17, 2007 contained three copepod species, viz., M. cf. gracilis, M. rubellus, and N. lacustris. The rain pool on Long Key sampled on November 1, 2007 contained Halicyclops sp. B and N. lacustris. It is not unexpected to find more than one species of copepod in a natural (i.e., not an artificial container) habitat (Fleeger 1985). As reviewed by Reid and Pinto-Coelho (1994), a wide range of vectors such as ship ballast, transport of fish and shellfish for aquaculture, and the aquarium trade have been implicated in the establishment of populations of non-native copepods. Ferrari and Rossetti (2006) discussed the finding of an Australasian copepod, Boeckella triarticulata (Thomson), in Italy; they suggested that the most likely vectors of introduction were stocking of allochthonous fish species and dispersal of resting eggs in imported crop seeds. Information from these and previous collections in southern Florida allows us to infer that the trade in ornamental plants may be an important vector in this region. The current collections bring the number of copepod species known from the Florida Keys to 33. As discussed by Reid and Hribar (2006), the assemblage of continental species includes nine Neotropical species (two calanoids and seven cyclopoids), six species found on more than one continent (five cyclopoids and one harpacticoid), and two North American cyclopoids; and is now known to include three exotic species, all of them cyclopoids. Most of the coastal marine-to-brackish-water species (three cyclopoids and ten harpacticoids) are widespread in the western Atlantic Ocean or worldwide, as far as their distribution is known. Southern Florida appears to harbor a moderately rich copepod fauna. However, the apparent species richness of an area oftentimes is a refl ection of the sampling effort expended in that area (Bruno et al. 2001). Doubtless further collecting will reveal additional species as yet undetected. 226 Southeastern Naturalist Vol.7, No. 2 Acknowledgments We thank Jody Davis, Emilio Posada, Lewis Robinson, and Carol Samul for their assistance in making collections. 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