A LIST AND IDENTIFICATION KEY FOR THE FRESHWATER, FREE-LIVING COPEPODS OF FLORIDA (U.S.A.) M. Cristina Bruno, Janet W. Reid, and Sue A.

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1 JOURNAL OF CRUSTACEAN BIOLOGY, 25(3): , 2005 A LIST AND IDENTIFICATION KEY FOR THE FRESHWATER, FREE-LIVING COPEPODS OF FLORIDA (U.S.A.) M. Cristina Bruno, Janet W. Reid, and Sue A. Perry (MCB, corresponding author) Department of Environmental Sciences, University della Tuscia, Largo dell Università snc, 01100, Viterbo, Italy (mcbruno@unitus.it); (JWR) Virginia Museum of Natural History, 1001 Douglas Avenue, Martinsville, Virginia 24112, U.S.A.; (SAP) South Florida Natural Resources Center, Everglades National Park, State Road 9336, Homestead, Florida 33034, U.S.A. ABSTRACT We present an inventory of the free-living freshwater copepod crustaceans recorded from the state of Florida, U.S.A. The list is based on previously published information and on new data collected during recent research on surface- and groundwater-dwelling copepods in Everglades National Park, and a few collections in temporary and permanent surface waterbodies elsewhere in the state. We provide information on the ecology and taxonomy of some of the species, and a key for the identification of all the taxa. A total of 65 taxa of freeliving copepods: 9 calanoids, 41 cyclopoids, and 15 harpacticoids are now known from the peninsula. Forty-four of these are known from Everglades National Park and adjacent areas; we add four species to a previous checklist for this region. The historically more intensive sampling here has resulted in the discovery of five new named taxa and six that remain in open nomenclature, 10 of which have so far been found only in the Everglades. Of the species collected so far in central and northern Florida, two calanoids and one cyclopoid have been found only in the state so far, whereas all the others are widespread in North America and beyond. Among the predominant North American fauna is a small neotropical component consisting of one calanoid, six cyclopoids, and five harpacticoids. One cyclopoid species is considered to be introduced. The Florida peninsula lies within the coastal plain, with a maximum elevation of 120 m (Webb, 1990); the highest ridges were formed as coastal dunes, and all major rock formations anywhere near the surface are marine sediments. Freshwater habitats in Florida are numerous and diverse. Florida has about 7800 lakes, covering about 6% of the landscape (Brenner et al., 1990); most of them have maximum depths less than 5 m (Kenner, 1964). The major river systems of Florida are in the northern and western region, but the rest of the peninsula has many small, slowly flowing streams. There are more than 300 springs, mostly artesian, with 27 springs of first magnitude (Nordlie, 1990). Owing to the low elevation and to the humid subtropical climate with a cool dry season and a warm rainy season, marshes and forested wetlands are a major component of the Florida landscape. More than half of the state was originally covered by wetlands (Shaw and Fredine, 1956); one-third of these freshwaters have disappeared (Hefner, 1986). Several large marshes are associated with the Kissimmee and St. John s Rivers; smaller marshes are scattered throughout the peninsula (Kushlan, 1990). The largest is the Everglades of southern Florida, a subtropical wetland ecosystem formed during the past 5000 years when prolonged inundation resulted in peat and marl accretion within the pre-existing limestone depression (see Bruno and Perry, 2004, and Bruno et al., 2003, for more physiographic and historical details). Everglades National Park (ENP) is a 0.9 million-hectare wilderness at the southern tip of Florida, bordered by Big Cypress National Preserve to the west and north, by Water Conservation Areas 3A and 3B to the north, and by developed urban and agricultural areas to the east along the Atlantic Coastal Ridge (Fig. 1). The Park includes part of the mangrove-lined coasts of Florida Bay, but is mostly occupied by freshwater sawgrass marshes interspersed with wet prairies, sloughs, and tree islands (hammocks). The Rocky Glades in eastern ENP contain marl prairies and surface bedrock, forming a typical karst landscape with thousands of solution holes (Hoffmeister, 1974). The freshwater planktonic fauna of central and northern Florida has been reported in numerous articles (e.g., Cowell et al., 1975; Bays and Crisman, 1983; Elmore et al., 1984). We have not attempted to provide a complete bibliography of this extensive literature, which of course mostly reports the most common planktonic species; rather, we have selected articles that provide as complete a list of species and county records as possible. In most of these studies, only lakes and ponds were sampled, usually over short periods. Only a few subterranean habitats have been investigated. The hyporheos, karst, and crenal of northern Florida have not been investigated, except for one sample series in interstitial bed sediments of the Blackwater River (Reid and Strayer, 1994), and another sample in Split Sink Cave in Wakulla County, from which the copepods recovered are listed here for the first time (Bowman and Sket, 1985, reported the isopod Remasellus parvus from the same collection). In only a few instances were copepods collected from more cryptic habitats in upper Florida, for instance ditches (Dickinson, 1949), a swamp (White et al., 1994; Leech and Wyngaard, 1996), leaf litter (Avery and Undeen, 1990), orchid roots (Lehman and Reid, 1992), and soil in a plant nursery (Reid, 1999). Bruno et al. (2003) inventoried the free-living copepod crustaceans of Everglades National Park and adjacent areas, providing detailed information on the ecology of the species and a taxonomic key for their identification. Further studies by MCB on planktonic copepod communities along the eastern border of ENP and in groundwater in the Rocky 384

2 BRUNO ET AL.: LIST AND IDENTIFICATION KEY FOR COPEPODS OF FLORIDA 385 Fig. 1. Extreme southern Florida, showing the boundaries of Everglades National Park, Big Cypress National Preserve, and Water Conservation Areas 3A and 3B. Collection sites (in circles) as coded in Table 2. Abbreviations: SRS, Shark River Slough; NESRS, Northeast Shark River Slough; TS, Taylor Slough; RG, Rocky Glades; WCA-3A, Water Conservation Area 3A; WCA-3B, Water Conservation Area 3B. Dotted areas: SRS, TS, NESRS. The inset map of the entire state shows the general locations where copepods have been collected. Glades have added a few more taxa to this list. We now present an inventory of all of the free-living copepods known from the entire state, based on published reports on surfaceand groundwater-dwelling copepods, the new records by MCB from the Everglades region, and additional, previously unpublished records from surface habitats in central and northern Florida. We list the individual locality records, with more detail for the new records. In a second table, we provide an updated checklist for the copepods known from the Everglades region, and summarize new and previously published information on collection sites, hydroperiod, and general ecological preferences. Finally, we present a key to aid in the identification of all the species and subspecies. MATERIALS AND METHODS Most of the records from ENP were reviewed and discussed by Bruno et al. (2003). Several new records were derived from long-term studies by MCB on surface- and groundwater communities. For the remaining Florida peninsula, we drew from published records (in several of these the original identifications were done by JWR), our own collections, and collections of other researchers examined by MCB or JWR.

3 386 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 25, NO. 3, 2005 For ENP and adjacent regions, almost all of the sites were described in detail by Bruno et al. (2003). The new additions are from three other studies. The first study consisted of monthly collections from surface waterbodies made during the wet seasons of 2002 and 2003 by MCB in retention ponds S-332B, S-332C, and S-332D on the eastern boundary of ENP. The retention ponds are a series of shallow infiltration basins constructed by the U.S. Army Corps of Engineers in to store water during the wet season for flood protection. From the detention areas, water seeps west into ENP and east into the canal system at a high rate because of the porous nature of the underlying limestone of the Biscayne Aquifer. The collections were made along three east-west transects of three stations in each detention area, plus one eastwest transect of three stations at Chekika. On each transect, one station was located within the detention area, on its western side, and two stations were located in ENP, approximately100 m and 500 m from the detention area. The second study consisted of surface-water samples collected by Dr. J. Trexler, Florida International University (FIU), which were examined by MCB. The samples were collected with funnel traps in three flumes constructed in the wet-prairie marshes of Shark River Slough. Each flume consisted of four channels used for P dosing at three different concentrations (see Noe et al., 2002, for details). The samples studied by MCB were collected on October 2003, representing the last series after five years of P dosing. They are stored in Dr. Trexler s laboratory at FIU. In a third study, to determine the spatial scale and distribution of the floating periphyton mat and its associated macroinvertebrate community, Smith and Trexler (2003) collected several samples in Shark River Slough and Water Conservation Area 3B. At each site, approximately cm diameter cores were taken from the periphyton mat, or the water column was sampled if a periphyton mat was not present. All copepods over 1 mm long were removed and given to MCB for identification. The specimens from this study are also stored in Dr. Trexler s laboratory at FIU. Additional new data from Bruno s studies in ENP include collections from crayfish holes made at the end of the dry season (May 2000), from surface water in Pine Glades Lake and Long Pine Key Campground Lake (April 2004), and two sets of wells in the Rocky Glades (Bruno and Perry, 2005). These samples are stored at the ENP South Florida Natural Resource Center Laboratory in Homestead. For ENP, sampling methods were reported in detail by Loftus and Reid (2000) and Bruno and co-workers (2000, 2001, 2002a, 2002b). For groundwater, we used a portable electric pump connected to a portable generator. For surface water, we used a hand pump for solution holes and crayfish holes, and inverted funnel traps for flooded areas. We also add new records from collections from a variety of surface waterbodies in southeastern Florida, made in 1995 by JWR and Willis A. Reid, Jr. Copepods were collected with a small plankton net, or from samples of substrate (wet leaves, moss, etc.). Several other, previously unpublished records are from material sent to JWR from various sources, as listed in Table 1. Samples of plankton collected in 1998 along the Kissimmee River by staff of the Riverwoods Field Laboratory were given to MCB for identification; the copepod specimens identified from these samples are stored at the ENP South Florida Natural Resource Center Laboratory in Homestead. There are a few samples of calanoid and cyclopoid copepods held in the collections of the National Museum of Natural History (Smithsonian Institution), which were identified by the eminent copepodologists Mildred S. Wilson and Thomas E. Bowman. As far as we can deterimne, these records were never published, and we have included them here. The new data are mostly from surface water, temporary (ponds, ditches, pools, soil, a storm drain) and some permanent (lakes, rivers) habitats in southeastern and central-eastern Florida. The few sites in the northern part of the state are also mostly from small surface waters, except for a few species collected by Boris Sket from a spring cave. The northwestern panhandle and the Florida Keys are nearly uninvestigated. Most of the localities in published reports are lakes (58 lakes in 22 counties); five are streams, creeks and rivers in five counties, five are wetlands in five counties, five are collections from soil or leaf litter in five counties, several are ponds and ditches in 10 counties, one site is a cave system, and one site is a beach (Table 1). Abbreviations used in the text: A1 ¼ antennule; A2 ¼ antenna; bsp ¼ basipodite; benp ¼ baseoendopodite; enp ¼ endopodite; enp 1 3, endopodite segments 1 3; exp ¼ exopodite; P1 5 ¼ thoracic legs 1 5. RESULTS A total of 65 taxa of free-living copepods: 9 calanoids, 41 cyclopoids, and 15 harpacticoids, are now known from the peninsula (Table 1). Forty-four of these are known from Everglades National Park and adjacent areas (Table 2). We add four species of cyclopoids (Ectocyclops sp., Eucyclops agilis, Microcyclops dubitabilis, and Paracyclops canadensis) to a previous checklist for ENP (Bruno et al., 2003). The historically more intensive sampling in ENP and environs has resulted in the discovery of five new, named taxa: Eucyclops conrowae, Thermocyclops parvus, Nitokra evergladensis, Elaphoidella fluviusherbae, and Elaphoidella marjoryae. Six additional species remain in open nomenclature: Ectocyclops sp., Metacyclops sp., Metacyclops cf. gracilis, Ectinosoma sp., Paramphiascella sp., and Parastenocaris sp. Except for E. conrowae, which has been found in several locations in the eastern U.S.A., all of these taxa have so far been found only in the Everglades. Of the species collected so far in central and northern Florida, the calanoids Aglaodiaptomus marshianus and Skistodiaptomus sinuatus, and the cyclopoid Diacyclops dimorphus have been found only in the state so far, whereas all the others are widespread in North America and beyond. Among the predominant North American fauna is a small neotropical component consisting of Arctodiaptomus dorsalis, Eucyclops bondi, Mesocyclops longisetus, Mesocyclops reidae, Metacyclops cushae, Microcyclops dubitabilis, Thermocyclops parvus, Nitokra bisetosa, N. evergladensis, E. fluviusherbae, E. marjoryae, and Elaphoidella grandidieri (which is pantropical). Bryocyclops muscicola is considered to be introduced (Reid, 1999). The individual records are dominated by the plankton of Central Florida lakes. This is mainly composed of three species of diaptomid calanoids, Arctodiaptomus dorsalis, Arctodiaptomus floridanus, and Skistodiaptomus mississippiensis, and two cyclopoids, Mesocyclops edax and Tropocyclops prasinus. One, or probably two species of cyclopoids, Ergasilus chautauquaensis and Ergasilus sp., the adult females of which are parasitic on fish, are also common (Table 1). Several species of littoral and benthic cyclopoids have been reported from permanent or temporary lentic habitats in central and northern Florida. Most of the reports refer to Acanthocylops vernalis, Microcyclops varicans, or Eucyclops agilis. Unfortunately, all these taxa are in need of revision. Other cyclopoids are reported less often (Table 1). The Kissimmee River is one of the few riverine habitats sampled so far, and information on its fauna is extremely sparse. Orthocyclops modestus was reported (as Orthocyclops) by Merritt et al. (1996). The plankton samples from the Kissimmee River identified by MCB contained S. mississippiensis, Macrocyclops albidus, and Eucyclops elegans. Records of species from habitats other than surface waters are scarce. Lehman and Reid (1992) reported Phyllognathopus viguieri from orchid roots in a nursery near Boynton Beach. Avery and Undeen (1990) cultured Microcyclops varicans from leaf litter collected in a swamp near Gainesville. Reid and Strayer (1994) described Diacyclops dimorphus, and later reported several other cyclopoid species (Strayer and Reid, 1999) from interstitial waters of the Blackwater River in northwestern Florida. Reid (1999) reported Bryocyclops muscicola from plant roots in a nursery in central Florida, and the same species was later found by

4 BRUNO ET AL.: LIST AND IDENTIFICATION KEY FOR COPEPODS OF FLORIDA 387 Table 1. List of copepod taxa reported in published records, and collections newly reported herein, for fresh waters of the state of Florida. ENP, Everglades National Park and adjacent areas; UFL, University of Florida, Gainesville. For previously unpublished records, institutions where specimens are deposited: ENP laboratory, South Florida Natural Resource Center Laboratory, Homestead; ENP Museum, Everglades National Park Museum, Homestead; USNM, National Museum of Natural History (Smithsonian Institution), Washington; VMNH, Virginia Museum of Natural History, Martinsville. Taxon Collection site Reference or collection data Order Calanoida Family Centropagidae Osphranticum labronectum S. A. Forbes, 1882 Family Diaptomidae Aglaodiaptomus clavipoides M. S. Wilson, 1955 Aglaodiaptomus marshianus M. S. Wilson, 1953 Arctodiaptomus dorsalis (Marsh, 1907) Arctodiaptomus floridanus (Marsh, 1926) ENP (a), see Table 2; Bear Bay Swamp, near Gainesville (Dixie County) (b); shallow slough in Big Cypress National Preserve (Monroe County) (c); roadside ditch near St. Marks (Wakulla County) (d). Unspecified (a). Lake Jackson (the type locality) and a nearby ditch pond (Leon County) (*a). Lakes Annie, Francis, Placid (Highland County), Blue Cypress (Indian River County), Eustis (Lake County), Geneva, Kingsley (Clay County), Jackson, Kerr, Weir (Marion County), Kingsley, Miona (Sumter County), Newnans, Wauberg (Alachua County), Ocean Pond (Baker County), Sampson, Santa Fe (Bradford County), Scott (Polk County), Washington (Brevard County) (a); Lake Okeechobee (Palm Beach County) (b, c, d, *e,*n); Tohopekaliga Lake (Osceola County) (a, f*); Lakes East Tohopegalika (Osceola County), Fells Cove (Osceola County), Kissimee (Osceola County), Cypress (Osceola County), Hatchineha (Osceola County) (*f); Lake Beauclair (Lake County) (g); Lake Thonotosassa (Hillsborough County), (h, i, j, *o, *p, *q, *r); East Lake (Seminole County) (j); Banana Lake (Polk County) (k); several ponds and ditches in Paynes Prairie State Park (Alachua County) (l); Lake Ruby (Orange County) (m); Rex Beach (Okaloosa County) (*n); Fairy Lake (Hillsborough County) (*p, *r); Little Lake George (Putnam County), Lake Monroe (Volusia County), St. Johns River (Brevard County) (*s); Middle Lake (Pasco County) (*s, w); small unnamed ponds in Indian River, Martin, and Palm Beach counties (t); Wakulla River (Wakulla County) (u); pond in Fort Pickens State Park (Escambia County) (v, w); Split Sink Cave system (Wakulla County) (x). ENP (a), see Table 2; Lake Annie (b, i); Lakes Blue Cypress, Eustis, Francis, Geneva, Kerr, Kingsley, Miona, Newnans, Ocean Pond, Placid, Sampson, Santa Fe, Scott, Tohopekaliga, Washington, Wauberg, Weir (b); Lake Jackson (b, **o); Corner and Bay lakes (Orange County) (c); Lake Thonotosassa (a, d); Spring Lake (Orange County) (e); Fairy Lake (f, g); grass carp ponds on UFL campus (Alachua County) (h); Lakes Ellen (Seminole County), Pretty (Hillsborough County), Padgett (Pasco County) (g); Lakes Altho (Alachua County), Anderson-Cue, Cowpen, Galilee, Rosa (Putnam County), Brooklyn, Magnolia, Sheeler (Clay County), Clay, Josephine, June), Kingsley, Francis, Letta (Highlands County), Geneva, Johnson, Lowery, McCloud (Polk County), Placid (i); Bivens Arm (Alachua County) and Newnans lakes (j); Lake Mize (j, k); Lake Ruby (l); Little Lake Conway, Lake Conway, Lake Gatlin (Orange County) (m); lakes near Lake City (Columbia County) (*n); pools in Polk County (the type locality) (*p); pond 3 miles (a) Bruno et al., 2003; (b) leg. S. E. White, 20 Jan 1994, det. JWR (USNM); (c) leg. JWR and W. A. Reid, Jr., 22 May 1995 (ENP Museum); (d) leg. I. Boliek, 8 Mar 1953, det. M. S. Wilson (USNM). (a) Wilson, May refer to specimens in the USNM collection, from a roadside pond north of Woodville (Leon County), leg. I. Boliek, 24 Apr 1955, det. M. S. Wilson. (a) Wilson, * as Diaptomus (Aglaodiaptomus) marshianus. (a) Bays and Crisman, 1983; (b) Crisman et al., 1995; (c) Carney et al., 1997; (d) Havens et al., 1996; (e) Havens and East, 1997; (f) Havens et al., 2000; (g) Elmore et al., 1984; (h) Elmore, 1983; (i) Elmore, 1982; (j) Dawes et al., 1987; (k) Osborne and Egan, 1997; (l) Dickinson, 1949; (m) Osborne and Jansen, 1993; (n) Kincaid, 1953; (o) Wyngaard, 1983; (p) Wyngaard, 1986a; (q) Wyngaard, 1986b; (r) Wyngaard, 1991; (s) Marsh, 1929; (t) coll. JWR and W. A. Reid, Jr., May 1995, det. JWR (ENP Museum); (*u) coll. I. Boliek, 16 Dec 1956, det. M. S. Wilson (USNM); (v) leg. G. E. Walsh, 15 Apr 1958, det. T. E. Bowman (USNM); (w) Suárez-Morales and Elías-Gutiérrez, 2001; (x) leg. B. Sket, Jul 1981, det. JWR (USNM; Split Sink system described by Bowman and Sket, 1985). * as Diaptomus dorsalis. (a) Bruno et al., 2003; (b) Bays and Crisman, 1983; (c) Elmore et al., 1984; (d) Cowell et al., 1975; (e) Billets and Osborne, 1985; (f) Elmore, 1983; (g) Dawes et al., 1987; (h) Fry and Osborne, 1980; (i) Brezonik et al., 1984; (j) Nordlie, 1976; (k) Harkness and Pierce, 1941; (l) Osborne and Jansen, 1993; (m) Blancher, 1984; (n) Kincaid, 1953; (o) Wilson, 1953; (p) Marsh, 1929; (*q) leg. W. G. Moore, 3 Sep 1954, det. M. S. Wilson (USNM), (r) Havens et al., * as Diaptomus floridanus. ** as Diaptomus (Arctodiaptomus) floridanus.

5 388 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 25, NO. 3, 2005 Table 1. Continued. Taxon Collection site Reference or collection data (4.8 km) southwest of Gainesville (Alachua County) (*q); Lakes Tohopekaliga; East Tohopegalika, Fells Cove, Kissimee, Cypress, Hatchineha (r*). Leptodiaptomus moorei M. S. Wilson, 1954 Unspecified (a). (a) Wilson, No specimens of this species from Florida are held in the USNM. Onychodiaptomus louisianensis M. S. Wilson and Moore, 1953 Skistodiaptomus mississippiensis (Marsh, 1894) Skistodiaptomus sinuatus Kincaid, 1953 Unspecified (a). Lakes Annie, Blue Cypress, Eustis, Francis, Geneva, Kerr, Kingsley, Miona, Newnans, Ocean Pond, Placid, Sampson, Santa Fe, Scott, Tohopekaliga, Washington, Wauberg, Weir (a); Lake Jackson (a, ** h); Corner and Bay lakes (Orange County) (b); Spring Lake (c); Fairy Lake (d, e); Lakes Ellen, Pretty, Padgett (e); several ponds and ditches in Paynes Prairie State Park (f); Kissimmee River (g); Lake Maitland (Orange County) (* i); Dog Pond (Leon County) (**j). Unnamed waterbody near Panama City (the type locality; Bay County) (*a); area of dry ponds, Florida University West Campus, Tallahassee (Leon County) (**b). Order Cyclopoida Family Cyclopidae Subfamily Eucyclopinae Ectocyclops sp. ENP, see Table 2. Ectocyclops phaleratus (Koch, 1838) ENP, see Table 2 (a); pond in Paynes Prairie State Park (b); pond in South Miami (Dade County) (c). Eucyclops agilis (Koch, 1838) ENP (a), see Table 2; Lake Thonotosassa (b); Spring Lake (c); grass carp ponds on UFL campus (d); pond in Paynes Prairie State Park (*e); headwater streams of the Alafia River (Hillsborough County) (f); pond in Boca Raton (Palm Beach County) and ditch in orange grove (Indian River County) (g); temporary woodland pool in Vero Beach (Indian River County) (h). Eucyclops agilis montanus (Brady, 1878) East Lake, Lakes Thonotosassa, Padgett (a). (a) Dawes et al., Eucyclops bondi Kiefer, 1934 ENP (a), see Table 2. (a) Bruno et al., Eucyclops conrowae Reid, 1992 ENP (a), see Table 2; slough in Big Cypress National Preserve, small constructed ponds in Deerfield Beach (Broward County), Linkport (Indian River County), and near Hobe Sound (Martin County), Blue Cypress Lake (b). Eucyclops elegans (Herrick, 1884) ENP (a), see Table 2; grass carp ponds on UFL campus (b); Kissimmee River (c); small constructed pond in Juno Beach (Palm Beach County) and blackwater creek near Dickinson State Park (Martin County) (d); Blue Heron Constructed Wetland, Titusville (Brevard County) (e). (a) Wilson, Wilson s (1959) record from Florida may refer to specimens in the USNM collection, from a roadside pond in Woodville (Leon County), leg. I. Boliek, 6 Dec 1956, det. M. S. Wilson. (a) Bays and Crisman, 1983; (b) Elmore et al., 1984; (c) Billets and Osborne, 1985; (d) Elmore, 1983; (e) Dawes et al., 1984; (f) Dickinson, 1949; (g) leg. C. de la Rosa, 1998, det. MCB (ENP laboratory); (h) Wilson, 1953; (i) Schacht, 1897; (j) leg. I. Boliek, 19 Mar 1957, det. M. S. Wilson (USNM). * as Diaptomus (Skistodiaptomus) mississippiensis. ** as Diaptomus mississippiensis. (a) Kincaid, 1953; (b) leg. I. Boliek, 22 Jul 1951, det. M. S. Wilson (USNM). * as Diaptomus (Skistodiaptomus) sinuatus. ** as Diaptomus sinuatus. (a) Bruno et al., 2003; (b) Dickinson, 1949; (c) leg. JWR and W. A. Reid, Jr., 23 May 1995, det. JWR (ENP Museum). (a) Bruno et al., 2003; (b) Cowell et al., 1975; (c) Billets and Osborne, 1985; (d) Fry and Osborne, 1980; (e) Dickinson, 1949; (f) Cowell et al., 2004; (g) leg. JWR and W. A. Reid, Jr., 23 and 28 May 1995, det. JWR (ENP Museum); (h) leg. J. R. Rey and S. O Connell, Feb 2004, det. JWR (VMNH). * as Cyclops serrulatus. (a) Bruno et al., 2003; (b) leg. JWR and W. A. Reid, Jr., May 1995, det. JWR (ENP Museum). (a) Bruno et al., 2003; (b) Fry and Osborne, 1980; (c) leg. C. de la Rosa, 2 October 1998, det. MCB (ENP laboratory); (d) leg. JWR and W. A. Reid, Jr., 24 May 1995, det. JWR (ENP Museum); (e) leg. J. R. Rey and S. O Connell, Apr 2004, det. JWR (VMNH). Eucyclops macrurus (G. O. Sars, 1863) Grass carp ponds on UFL campus (a). (a) Fry and Osborne, Homocyclops ater (Herrick, 1882) ENP (a), see Table 2; several ponds and ditches in (a) Bruno et al., 2003; (b) Dickinson, Paynes Prairie State Park (b). Macrocyclops albidus (Jurine, 1820) ENP (a), see Table 2; Spring Lake (b): several ponds and ditches in Paynes Prairie State Park (a) Bruno et al., 2003; (b) Billets and Osborne, 1985; (c) Dickinson, 1949; (d) leg. C. de la (c); Kissimmee River (d); Bear Bay Swamp (e); Rosa, 2 October 1998, det. MCB (ENP headwater streams of the Alafia River (f); slough in Big Cypress National Preserve, small constructed ponds and ditches in Dade, Indian River, Martin, and Palm Beach counties, and Blue Cypress Lake (g); cultures, Panama City (see text) (h); temporary woodland pool in Vero Beach (i); Split Sink Cave system (j). laboratory); (e) leg. S. E. White, 20 Jan 1994, det. JWR (USNM); (f) Cowell et al., 2004; (g) leg. JWR and W. A. Reid, Jr., May 1995, det. JWR (ENP Museum); (h) Rey et al., 2004; (i) leg J. R. Rey and S. O Connell, Feb 2004, det. JWR (specimens discarded); (j) leg. B. Sket, Jul 1981, det. JWR (USNM; Split Sink system described by Bowman and Sket, 1985).

6 BRUNO ET AL.: LIST AND IDENTIFICATION KEY FOR COPEPODS OF FLORIDA 389 Table 1. Continued. Taxon Collection site Reference or collection data Macrocyclops fuscus (Jurine, 1820) ENP (a), see Table 2. (a) Bruno et al., Paracyclops canadensis (Willey, 1934) ENP, see Table 2. Paracyclops chiltoni (Thomson, 1883) ENP (a), see Table 2; Banana Lake (b); Lake Ruby (c); Bear Bay Swamp (*d); Blackwater River (Santa Rosa County) (e). (a) Bruno et al., 2003; (b) Osborne and Egan, 1997 (as P. fimbriatus); (c) Osborne and Jansen, 1993 (as P. fimbriatus); (d) White et al., 1994); (e) Strayer and Reid, * as P. fimbriatus chiltoni Paracyclops poppei (Rehberg, 1880) ENP (a), see Table 2; Fairy Lake (b); headwater streams of Alafia River (*c). (a) Bruno et al., 2003; (b) Dawes et al., 1987; (c) Cowell et al., * as P. fimbriatus poppei Paracyclops sp. (undescribed) Blackwater River (a). (a) Strayer and Reid, Tropocyclops extensus Kiefer, 1931 ENP (a), see Table 2; Blue Heron Constructed (a) Bruno et al., 2003; (b) leg. J. R. Rey and S. Wetland, Titusville (b). O Connell, Apr 2004, det. JWR (VMNH). Tropocyclops prasinus (Fischer, 1860) Lake Okeechobee (a, b, c, d, e); Lakes Tohopekaliga; East Tohopegalika, Fells Cove, Kissimee, Cypress, Hatchineha (f); Lake Beauclair, Corner, and Bay lakes (g); Spring Lake (h); grass carp ponds on UFL campus (i); Lakes East, Ellen, Fairy, Padgett, Pretty, Thonotosassa (j), Banana Lake (k); Lakes Anderson-Cue, (a) Crisman et al., 1995; (b) Carney et al., 1997; (c) Havens et al., 1996; (d) Havens and Beaver, 1997; (e) Havens and East, 1997, (f) Havens et al., 2000; (g) Elmore et al., 1984; (h) Billets and Osborne, 1985; (i) Fry and Osborne, 1980; (ji) Dawes et al., 1987 (k) Osborne and Egan, 1997; (l) Brezonik et al., 1984; (m) Nordlie, Annie, Clay, Cowpen, Francis, Galilee, Geneva, 1976; (n) Osborne and Jansen, 1993; (o) Johnson, Josephine, June, Kingsley, Letta, Blancher, 1984; (p) Forbes, 1897; (q) leg. Lowery, McCloud, Rosa, Placid, Sheeler, (l); Lake Mize (m); Lake Ruby (n); Little Lake JWR and W. A. Reid, Jr., 24 May 1995, det. JWR (ENP Museum). * as Cyclops prasinus. Conway, Lakes Conway and Gatlin (o); Sister Lake (Charlotte County) (*p); small constructed pond in Juno Beach (Palm Beach County) (q). Tropocyclops prasinus mexicanus Kiefer, 1938 Subfamily Cyclopinae Acanthocyclops robustus (G. O. Sars, 1863) s.l. Acanthocyclops vernalis (Fischer, 1853) Bryocyclops muscicola (Menzel, 1926) ENP (a), see Table 2; small constructed ponds in South Miami (Dade County) and Hobe Sound (Martin County), blackwater creek near Dickinson State Park (Martin County), and Blue Cypress Lake (b). ENP (a), see Table 2; ponds and ditches in Indian River, Martin, and Palm Beach counties (b). Lake Okeechobee (a, b, c, d, e); Lakes Tohopekaliga; East Tohopegalika, Fells Cove, Kissimee, Cypress, Hatchineha (f); Lake Beauclair (*g); East Lake, Lakes Ellen, Pretty, Thonotosassa (h); Little Lake Conway, Lakes Conway and Gatlin (i). Soil in plant nursery, Zellwood (Orange County) (a); soil in plant nursery, Apopka (Lake County) (b). (a) Bruno et al., 2003; (b) leg. JWR and W. A. Reid, Jr., May 1995, det. JWR (ENP Museum). (a) Bruno et al., 2003; (b) leg. JWR and W. A. Reid, Jr., May 1995, det. JWR (ENP Museum). (a) Crisman et al., 1995; (b) Carney et al., 1997; (c) Havens et al., 1996; (d) Havens and Beaver, 1997; (e) Havens and east, 1997; (f) Havens et al., 2000; (g) Elmore et al., 1984; (h) Dawes et al., 1987; (i) Blancher, * as Cyclops vernalis. (a) Reid, 1999; (b) leg. P. S. Lehman, 1997, det. JWR (USNM). [Populations considered to be introduced.] Diacyclops crassicaudis brachycercus ENP (a), see Table 2. (a) Bruno et al., (Kiefer, 1927) Diacyclops dimorphus Reid et Strayer, 1994 Blackwater River (a, b). (a) Reid and Strayer, 1994; (b) Strayer and Reid, Diacyclops navus (Herrick, 1882) Bear Bay Swamp (a, b). (a) Leech and Wyngaard, 1996; (b) Dorward and Wyngaard, Diacyclops nearcticus Kiefer, 1934 ENP (a), see Table 2; Bear Bay Swamp (b). (a) Bruno et al., 2003; (b) leg. S. E. White, 17 Nov 1993, det. JWR (USNM). Diacyclops sororum Reid, 1992 Blackwater River (a). (a) Strayer and Reid, Diacyclops thomasi (S. A. Forbes, 1882) Lake Thonotosassa (a). (a) Cowell et al., Mesocyclops americanus Dussart, 1985 ENP (a), see Table 2; Lake Mize (*b). (a) Bruno et al., 2003; (b) Harkness and Pierce, * as Cyclops leuckarti. Mesocyclops edax (S. A. Forbes, 1891) ENP (a), see Table 2; Lake Okeechobee (b, c, d, e, f); Lakes Tohopekaliga; East Tohopegalika, Fells Cove, Kissimee, Cypress, Hatchineha (g); Lake Beauclair, Bay and Corner lakes (h); Lake Thonotosassa (i, j, k, l, v, w, x, y, z); Fairy Lake (j, l, w, x, y), East Lake, Lakes Ellen, Pretty, Padgett (l); Spring Lake (m); grass carp ponds on UFL campus (n); Banana Lake (o); Lakes Altho, Anderson-Cue, Annie, Brooklyn, Clay, Cowpen, Francis Galilee, Geneva, Johnson, Josephine, June, Kingsley, Letta, Lowery, Magnolia, McCloud, Placid, Rosa, Sheeler (p); Bivens Arm and Newnans lakes, Lake Mize (q); (a) Bruno et al., 2003; (b) Crisman et al., 1995; (c) Carney et al., 1997; (d) Havens et al., 1996; (e) Havens and East, 1997; (f) Havens and Beaver, 1997; (g) Havens et al., 2000; (h) Elmore et al., 1984; (i) Cowell et al., 1975; (j) Wyngaard et al., 1985; (k) Wyngaard et al., 1982; (l) Dawes et al., 1987; (m) Billets and Osborne, 1985; (n) Fry and Osborne, 1980; (o) Osborne and Egan, 1997; (p) Brezonik et al., 1984; (q) Nordlie, 1976; (r) Osborne and Jansen, 1993; (sr) Blancher, 1984; (t) leg. C. de la Rosa, 2 October 1998, det. MCB (ENP laboratory); (u) E. B. Forbes, 1897; (v) Wyngaard, 1983; (w)

7 390 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 25, NO. 3, 2005 Table 1. Continued. Taxon Collection site Reference or collection data Lake Ruby (r); Little Lake Conway, Lakes Conway and Gatlin (s); Kissimmee River (t); Sister Lake, Lake Butler (Orange County) (*u); Split Sink Cave system (1). Wyngaard, 1986a; (x) Wyngaard, 1986b; (y) Wyngaard, 1991; (z) Wyngaard et al., 1994; (1) leg. B. Sket, Jul 1981, det. JWR (USNM; Split Sink system described by Bowman and Sket, 1985). * as Cyclops edax. Mesocyclops longisetus (Thiébaud, 1912) Florida, unspecified locality (a); central Florida (b). (a) Dussart and Fernando, 1986; (b) Tietze et al., Mesocyclops reidae Petkovski, 1986 ENP (a), see Table 2. (a) Bruno et al., Metacyclops cushae Reid, 1991 ENP (a), see Table 2. (a) Bruno et al., Metacyclops sp. ENP (a), see Table 2. (a) Bruno et al., Metacyclops cf. gracilis (Lilljeborg, 1853) ENP (a), see Table 2. (a) Bruno et al., Microcyclops rubellus (Lilljeborg, 1901) Microcyclops varicans (G. O. Sars, 1862) ENP (a), see Table 2; East Lake, Lakes Thonotosassa, Ellen (b); grass carp ponds on UFL campus (c); Bear Bay Swamp (d); (e) Shark River Slough in ENP, slough in Big Cypress National Preserve, and small constructed ponds in Linkport and Vero Beach, and ditch in orange grove (Indian River County). ENP (a), see Table 2; cultured from leaf litter in Gainesville (b); Banana Lake (c); Lakes Altho, Anderson-Cue, Annie, Brooklyn, Clay, Cowpen, Francis, Galilee, Geneva, Johnson, Josephine, June, Kingsley, Letta, Lowery, Magnolia, McCloud, Placid, Rosa, Sheeler, (d); storm drain catch basin, Key West (Monroe County) (e). (a) Bruno, 2003; (b) Dawes et al., 1987; (c) Fry and Osborne, 1980; (d) leg. S. E. White, 20 Jan 1994, det. JWR (USNM); (e) leg. JWR and W. A. Reid, Jr., May 1995, det. JWR (ENP Museum). (a) Bruno, 2003; (b) Avery and Undeen, 1990; (c) Osborne and Egan, 1997; (d) Brezonik et al., 1984; (e) leg. J. R. Rey and S. O Connell, Jun 2004, det. JWR (specimens discarded). Microcyclops dubitabilis (Kiefer, 1934) ENP, see Table 2. Orthocyclops modestus (Herrick, 1883) ENP (a), see Table 2; Kissimmee River (*b). (a) Bruno et al., 2003; Merritt et al., * as Orthocyclops. Thermocyclops parvus Reid, 1989 ENP (a), see Table 2. (a) Bruno et al., Family Ergasilidae Ergasilus chautauquaensis Fellows, 1887 Ergasilus sp. Lake Thonotosassa (a, b); Lakes East, Ellen, Fairy, (a) Cowell et al., 1975; (b) Dawes et al., Padgett, Pretty (b). Lakes Anderson-Cue, Annie, Brooklyn, Clay, (a) Brezonik et al., Galilee, Geneva, Johnson, Josephine, Letta, Lowery, Magnolia, McCloud, Rosa (a). Order Harpacticoida Family Ectinosomatidae Ectinosoma sp. ENP (a), see Table 2. (a) Bruno et al., Family Phyllognathopodidae Phyllognathopus viguieri (Maupas, 1892) ENP (a), see Table 2; orchid roots in nursery near Boynton Beach (Palm Beach County) (b); soil in plant nursery, Miami (Dade County) (c); soil in plant nursery, Zellwood (Orange County) (d). Family Diosaccidae Paramphiascella sp. ENP (a), see Table 2. (a) Bruno et al., Family Ameiridae Nitokra bisetosa Mielke, 1993 ENP (a), see Table 2. (a) Bruno et al., Nitokra evergladensis Bruno et Reid, 2002 ENP (a), see Table 2. (a) Bruno et al., Family Canthocamptidae Attheyella americana (Herrick, 1884) ENP (a), see Table 2; Lake Jackson (b); Bear Bay Swamp (c). (a) Bruno et al., 2003; (b) Lehman and Reid, 1992; (c) leg. P. S. Lehman, 1996, det. JWR (USNM); (d) leg. P. S. Lehman, 1997, det. JWR (USNM). (a) Bruno et al., 2003; (b) M. S. Wilson, 1958; (c) leg. S. E. White, 17 Nov 1993, det. JWR (USNM). Bryocamptus hutchinsoni Kiefer, 1929 Bear Bay Swamp (a). (a) leg. S. E. White, 29 Jan 1994, det. JWR (USNM). Bryocamptus newyorkensis (Chappuis, 1927) ENP (a), see Table 2. (a) Bruno et al., Canthocamptus sp. Grass carp ponds on UFL campus (a). (a) Fry and Osborne, Cletocamptus deitersi (Richard, 1897) ENP (a), see Table 2. (a) Bruno et al., Elaphoidella fluviusherbae ENP (a), see Table 2. (a) Bruno et al., Bruno and Reid, 2000 Elaphoidella marjoryae Bruno and Reid, 2000 ENP (a), see Table 2. (a) Bruno et al., Elaphoidella grandidieri (Guerne and Richard, 1893) ENP (a), see Table 2. (a) Bruno et al., 2003.

8 BRUNO ET AL.: LIST AND IDENTIFICATION KEY FOR COPEPODS OF FLORIDA 391 Table 1. Continued. Taxon Collection site Reference or collection data Family Parastenocarididae Parastenocaris sp. ENP (a), see Table 2. (a) Bruno et al., Family Laophontidae Onychocamptus mohammed (Blanchard and Richard, 1891) ENP (a), see Table 2. (a) Bruno et al., P. S. Lehman in another nursery (Table 1). Bryocyclops muscicola was previously known from Java and Sumatra, and the populations in Florida were likely introduced by human agency (Reid, 1999). Sket (as described by Bowman and Sket, 1985) made the only collection including copepods from a cave system of which we are aware. In 1993, adults of Acanthocyclops vernalis and Mesocyclops longisetus, from cultures originating in Louisiana, were released into outdoor tire piles as part of mosquito control experiments carried out in Panama City (Hallmon et al., 1993; Schreiber et al., 1993, 1996; Tietze et al., 1994). The original cultures were established by Dr. Gerald G. Marten, then of the New Orleans Mosquito Control Board, and the species were identified by JWR. It is not known whether these copepods ever established local populations outside the tires. We have listed this locality record in Table 1. Both of the species have been reported from elsewhere in Florida. For ENP, Table 2 gives a detailed list of the collection localities for the taxa reported by Bruno et al. (2003) and the new records. Most of the taxa reported by Bruno et al. (2003) and in works cited therein are not further discussed here. In the following several paragraphs we briefly discuss taxonomic or systematic problems with certain species. In most cases, we have not attempted to locate specimens from older studies in order to verify previous dubious records. In respect to taxonomic questions, we describe our own analyses of material. The first problematic group is the Acanthocyclops vernalis-robustus complex. In North America, this group contains several named and unnamed taxa, some of which have notoriously resisted morphological or genetic analysis. Past confusions have been much discussed (e.g., Dodson, 1994, and references therein). The morphological characters of these cyclopoids are unusually plastic to environmental factors, and in a recent study, Dodson et al. (2003) found little correlation between morphology and reproductive isolation as evaluated by crossbreeding experiments in the laboratory. In our samples from the Everglades, we found a single morph that corresponds roughly with the Acanthocyclops robustus warm-water strain of Dodson (1994). Because in the present state of understanding it is impossible to guess how many biological species of this group might be living in Florida, the published records listed in Table 1 must be treated with caution. Karanovic (2000) proposed a new genus Reidcyclops to include Diacyclops dimorphus and two species from Macedonia and the Caucasus. Reidcyclops was defined partly on the structure of leg 5. In the Florida species, this leg is typically diacyclopine, i.e., with a slender but definitely spiniform inner subterminal spine next to the outer terminal seta, rather than the two slender terminal setae borne by the European species. The structure of the swimming legs is also different in D. dimorphus, with fewer reductions in setae. The exopodites of both P3 and P4 are 3-segmented in males of the European forms; whereas in the Florida species only the P4 exopodite retains the 3-segmented structure. Therefore, we prefer to retain the taxon dimorphus in the genus Diacyclops. A few specimens listed as Ectocyclops sp., collected in ENP, have the antennule 11-segmented (in the females), and the middle seta of leg 5 shorter than the others. These characters are shared by E. polyspinosus Harada, 1931, which was described from Taiwan and reported from lakes in Ontario, Canada, by Dussart and Fernando (1990). However, because several nominal species of the genus are variable, as discussed for example by Morton (1990), we prefer not to assign a species name until more material becomes available. Eucyclops agilis, a much-employed synonym of which is E. serrulatus (Fischer, 1851), closely resembles E. conrowae. Several subtle differences between the two species were listed by Reid (1992). Eucyclops agilis is a common and possibly cosmopolitan species (Dussart and Defaye, 1985), and has been recorded in Florida several times (Table 1). It is likely that some of the older records refer to E. conrowae. Eucyclops speratus (Lilljeborg, 1901), reported by Fry and Osborne (1980), is an Eurasian species (Ishida and Hiruta, 1999). In North America, most of the populations of a Eucyclops with long caudal rami are referable to E. elegans, and we have accordingly listed the record under the latter name. Paracyclops chiltoni was long considered a subspecies of P. fimbriatus, until the recent revision by Karaytug (Karaytug and Boxshall, 1998b; Karaytug, 1999). We follow Karaytug s opinion that P. fimbriatus s.str. does not occur in North America, and accordingly have listed the records of this species under P. chiltoni in Table 1. Specimens of a species of Metacyclops that are similar but not identical to M. gracilis (Lilljeborg, 1853) were collected from groundwater in the Rocky Glades (Bruno and Perry, 2004), and one specimen was collected in a solution hole at the end of the dry season (Table 2). Metacyclops gracilis has been reported from Europe, Africa, Asia, and South America (Argentina) (Dussart and Defaye, 1985), but the few previous records of M. gracilis from North America are uncertain (Reid, 1991). There are few differences between the Everglades specimens and the description of M. gracilis. In M. gracilis, the length of the P4-enp3 outer terminal seta is only about one-fifth the length of the inner terminal seta (Gurney, 1933; Dussart, 1969), whereas in the Everglades specimens it is about one-third the length of the outer terminal seta. In the caudal ramus of M. gracilis, the innermost terminal caudal seta is about twice the length of

9 392 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 25, NO. 3, 2005 Table 2. Free-living copepod species in freshwater habitats of Everglades National Park and adjacent areas. Collection codes: 1, Reid (1992); 2, Reid (1989); 3, Loftus and Reid (2000); 4, Bruno et al. (2000); 5, Bruno et al. (2001); 6, Bruno et al. (2002b); 7, Bruno et al. (2002a); 8, Bruno et al., 2003; 9, Bruno and Perry, 2004; 10, Bruno and Perry, 2005; 11, retention areas transects (see text for details); 12, bromeliads, Anhinga Trail, ENP, October 2001; 13, crayfish holes and solution holes, Pa-hay-okee and Rocky Glades, March 2002; 14, S. L. Smith floc and periphyton mat collections in Shark River Slough, Northeast Shark River Slough, WCA3A (Smith and Trexler, 2003); 15, Pine Glades Lake and Long Pine Key Campground Lake, June 2004; 16, J. Trexler flumes collections in Shark River Slough (see text for details). Collection site codes as follows: A: Rocky Glades, surface water, wet season; B: Rocky Glades, solution holes, dry season; C: Rocky Glades, rehydrated soil, dry season; D: Rocky Glades, Long Pine Key wells; E: Taylor Slough, surface water, wet season; F: Taylor Slough, rehydrated soil, dry season; G: Pa-hay-okee, surface water, wet season; H, Pa-hay-okee, crayfish holes, dry season; I: Shark River Slough, surface water; J: Shark River Slough, rehydrated soil, dry season; K: Northeast Shark River Slough, surface water; L: Northeast Shark River Slough, rehydrated soil, dry season; M: Long Lake, surface water; N: north-central Dade County, wells; O: detention areas S-332B, C, D, surface water; P: Big Cypress, surface water; Q: bromeliad phytotelmata; R: east Everglades; S: Rocky Glades, Chekika; T: Water Conservation 3A; U; Pine Glades lake; V: Long Pine Key campground lake. Hydroperiod codes: s: short; i: intermediate; l: long; v: variable; p: permanent water; na: not applicable. Ecological characteristics: Ep: epigean; stx: stygoxene; stp: stygophile; stb: stygobite; br: bromeliads. Taxon Collection site, Hydroperiod A, s B, s C, s D, s E, i F, i G, i H, i I, l J, l K, I L, p M, l N, na O, v P, l Q, v R, i S, s T, l U, p V, p Order Calanoida Osphranticum labronectum 6 5 9, 10 6, , 14, 16 1, , Ep, stx Arctodiaptomus floridanus , , 11 Ep, stx Order Cyclopoida Acanthocyclops robustus , 10 6, , , Ep, stx Diacyclops crassicaudis brachycercus 8 Stp Diacyclops nearcticus 5 4, 9, Stp Ectocyclops sp Ep Ectocyclops phaleratus , 11 Ep, stx Eucyclops agilis Ep Eucyclops bondi Ep Eucyclops conrowae 6 9 6, , , Ep, stx Eucyclops elegans 6, , 11 Ep, stx Homocyclops ater 6 9 6, , 16 8 Ep, stx Macrocyclops albidus 6 5 9, 10 6, , 14, 16 1, , Ep, stx Macrocyclops fuscus Ep, stx Mesocyclops americanus 6 5 9, 10 6, , 11 Ep, stx Mesocyclops edax , Ep, stx Mesocyclops reidae 8 Ep, stx Metacyclops cushae 8 Ep, stx Metacyclops sp. 12 Br Metacyclops cf. gracilis 13 9 Stp? Microcyclops rubellus , 10 6, , 11 Ep, stx Microcyclops varicans 6 5 9, 10 6, , 11 Ep, stx Microcyclops dubitabilis Ep, stx Orthocyclops modestus 6 9, Ep, stp Paracyclops canadensis Ep Paracyclops chiltoni Ep, stx Paracyclops poppei , , 11 Ep, stx Tropocyclops extensus Stp? Thermocyclops parvus 6 5 9, 10 6, , , 11 Ep, stp Tropocyclops prasinus mexicanus 6 5 9, 10 6, , Ep, stx Order Harpacticoida Attheyella americana 7 6, 11 Ep Bryocamptus newyorkensis 7, 9 6, 7 5, 7 Ep Cletocamptus deitersi 1, Ep Ectinosoma sp. 9 Stx Elaphoidella fluviusherbae 5 4, 9 Stb Elaphoidella marjoryae 4, 9 Ep, stp Elaphoidella grandidieri 10, Ep Nitokra bisetosa 10 trap Stx Nitokra evergladensis 6, 7, Ep Onychocamptus mohammed , 16 1 Ep Paramphiascella sp. 10 trap Stx Parastenocaris sp. 9 Stb Phyllognathopus viguieri 1, Ep, stx Ecology the outermost terminal caudal seta (Gurney, 1933; Dussart, 1969), whereas their lengths are almost equal in the Everglades species. In M. gracilis, the A2 basipodite bears one seta (Gurney, 1933), whereas the Everglades specimens have two. However, in our opinion the morphological differences from the nominal species are insufficient to establish a new taxon. Collection of more specimens will be needed to check for variability and for further comparison

10 BRUNO ET AL.: LIST AND IDENTIFICATION KEY FOR COPEPODS OF FLORIDA 393 with M. gracilis. Unfortunately, M. cf. gracilis is rare in ENP and adapted to live in groundwater; it was not collected during the several previous studies of surface waters (Reid, 1989, 1994; Loftus and Reid, 2000; Bruno et al., 2001, 2002b), or of groundwater copepods in ENP (Bruno and Perry, 2004) or adjacent areas (Bruno et al., 2003). Several specimens of Microcyclops dubitabilis were collected from surface water in the detention areas east of ENP, as well as in the eastern part of the park. This species is reported from Mexico, Central and South America, and the Antilles (Reid, 1985, 1990; Dussart and Defaye, 1985; Löffler, 1981). This is the first report of this species in North America. According to Reid (1992), Everglades populations of M. varicans differ from populations of M. rubellus primarily in the short lateral terminal spine of the P4 endopodite terminal segment (medial spine/lateral spine length ratios: 1.9 in females of M. varicans, and 1.3 in females of M. rubellus), and by the few stout spines along the posterior margin of the anal somite (whereas M. rubellus has spines along the margin, increasing in size medially). Fiers et al. (2000), describing specimens of M. dubitabilis from a cenote (sinkhole) in the Yucatán, pointed out the morphological differences between this species and M. rubellus (as described in Reid, 1992, from specimens collected in Everglades National Park), and distinguished the two species on the basis of the length of the medial seta on the caudal ramus, and the number of rows of slender spinules on the caudal surface of the coxopodite of P4. According to Fiers et al. (2001), specific features of M. dubitabilis are the length/width ratio of the caudal ramus (females 2.6, males 2.25) the long inner apical seta on the caudal ramus, and the length of the medial spine of the P4 endopodite terminal segment relative to the length of the supporting segment (80% to 93% for both sexes); both species have the posterior margin of the anal somite entirely lined with spinules. Fiers et al. (2000) did not discuss the similarities with M. varicans, even though M. dubitabilis and M. varicans share the character of the P4 endopodite terminal spines of unequal lengths. Reid (1985) suggested that M. dubitabilis may be a synonym of M. varicans. However, further examination of our material from ENP showed that the three species have distinguishable features, and also that ENP populations vary slightly from the published descriptions in respect to the most easily distinguishable characters (the ornamentation of the posterior margin of the anal somite; the relative length of the P4 endopodite terminal spines; the ratio of the leg 4 medial terminal spine to the segment bearing it): 1. Microcyclops dubitabilis: Spines extending along the entire posterior margin of the anal somite in both sexes. Female antennule 12-segmented. The mean length/width ratio of the caudal ramus in both sexes (2.5 for females and 2.25 for males) in the Everglades specimens corresponds almost exactly to that given by Fiers et al. (2000) for M. dubitabilis, and also to the P4-enp2 medial terminal spine length (mean 78% of the P4-enp2 segment for females, and 87% for males). 2. Microcyclops rubellus: Spines along the posterior margin of the anal somite. As observed by Reid (1985), in the females of ENP populations, the spines are large and increase in size medially. We observed a group of 4 12 larger ventral spines and a row of very tiny spines, which are sometimes difficult to see under ordinary magnifications, extending laterally around the margin. In males, all of the spines appear to be the same size. Reid (1992) observed only females with 11-segmented antennules; however, we recorded a few females with antennules of 12 segments. 3. Microcyclops varicans: Spines along the posteroventral margin of the anal somite. Reid (1985) reported that in Florida populations of M. varicans, both sexes bear 4 6 spines along the central margin, not extending to the lateral margin of the somite; the spines are large and increase in size medially. Gurney (1933) also described the ornamentation in this species as a group of spines, often reduced to six spines, not extending to the sides. Observing many additional individuals, we found that the number of spines is more variable: the Everglades specimens may have either one group of 4 11 large spines, or two groups: a medial group of 3 9 larger spines, and a more lateral group of 3 8 smaller spines, for a maximum total of 14 spines, which all together do not reach past the side of the somite. The antennule is uniformly 12-segmented in females. Microcyclops varicans has the longest medial terminal spine on the P4enp2, relative to the supporting segment (spine length on average 85% of the segment in females). In females, the mean ratio of the medial terminal spine/lateral terminal spine on this segment is Paracyclops canadensis (Willey, 1934) was originally described from specimens collected in Québec, Canada. According to Karaytug (1999), all of the records of Paracyclops affinis (G. O. Sars, 1863) from North America actually refer to P. canadensis. This species is most often found in acid bogs, where it is often present in the leaf cups of the pitcher plant Sarracenia purpurea (Hamilton et al., 2000). Its known range extends from southeastern Canada through the central and eastern U.S.A. as far as South Carolina (Karaytug and Boxshall, 1998a; Karaytug, 1999; Hamilton et al., 2000). The find of several specimens in surface waters near the eastern border of ENP extends the distribution considerably southwards. The ability to occupy temporary habitats may have helped P. canadensis to colonize the Everglades ephemeral marshes. Tropocyclops extensus was described by Kiefer (1931) from specimens collected in New Jersey. Kiefer later (1934) gave some measurements (but no figures) of a single female from a collection in Haiti. This morph had long caudal rami (just over 3 times longer than wide), i.e., longer than in members of the Tropocyclops prasinus-group as it was then understood. Kiefer tentatively assigned the Haitian morph to the taxon Eucyclops (Tropocyclops) cf. extensus. The New Jersey population had considerably longer caudal rami (length: width ratio ). Dussart and Fernando (1990) gave some measurements for a form that they ascribed to T. extensus (which was collected in a bog in Ontario, Canada), if we consider that Kiefer s form is an extreme case considering that the furca is long relative to the total length (Dussart and Fernando, 1990: 2601). Bruno and Perry (2004) reported specimens of Tropocyclops, tentative-