Site by Bennett Web & Design Co.
2008 SOUTHEASTERN NATURALIST 7(2):219–228
New Records of Copepods (Crustacea) from the
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.
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 - firstname.lastname@example.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
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
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
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).
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.
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
We thank Jody Davis, Emilio Posada, Lewis Robinson, and Carol Samul for their
assistance in making collections. Two anonymous reviewers, Dr. Elena G. Krupa,
Dr. Eduardo Suárez-Morales, and Dr. Ricardo Pinto-Coelho made comments that
resulted in significant improvements to the manuscript.
Blumenshine, S.C., and K.D. Hambright. 2003. Top-down control in pelagic systems:
A role for invertebrate predation. Hydrobiologia 491:347–356.
Bonou, C.A., M. Pagano, and L. Saint-Jean. 1991. Développement et croissance
en poids de Moina (cf) micrura et de Mesocyclops ogunnus dans un milieu
saumâtre tropical: Les étangs de pisciculture de Layo (Côte-d’Ivoire). Revue
d’Hydrobiologie Tropicale 24:287–303.
Bruno, M.C., W.F. Loftus, and S.A. Perry. 2001. Preliminary data on microcrustacean
communities from ground waters in the southern Everglades. Pp. 89–97, In
E.L. Kuniansky (Ed.). US Geological Survey Karst Interest Group Proceedings,
Water-Resources Investigations Report 01–4011. 211 pp.
Bruno, M.C., J.W. Reid, and S.A. Perry. 2005. A list and identification key for the
freshwater, free-living copepods (Crustacea) of Florida (USA). Journal of Crustacean
Cathcart, D.J. 1995. The importance of maintaining bromeliad imports. Florida Entomologist
Confer, J.L. 1971. Intrazooplankton predation by Mesocyclops edax at natural prey
densities. Limnology and Oceanography 16:663–666.
Dobrzykowski, A.E., and G.A. Wyngaard. 1993. Phenology of dormancy in a Virginia
population of Mesocyclops edax (Crustacea: Copepoda). Hydrobiologia
Ferrari, I., and G. Rossetti. 2006. New records of the centropagid Boeckella
triarticulata (Thomson, 1883) (Copepoda: Calanoida) in northern Italy: Evidence
of a successful invasion? Aquatic Invasions 1:219–222.
Fleeger, J.W. 1985. Meiofaunal densities and copepod species composition in a
Louisiana, USA, estuary. Transactions of the American Microscopical Society
Gophen, M. 1976. Temperature effect on lifespan, metabolism, and development
time of Mesocyclops leuckarti (Claus). Oecologia 25:271–277.
Gophen, M. 1977. Food and feeding habits of Mesocyclops leuckarti (Claus) in Lake
Kinneret (Israel). Freshwater Biology 7:513–518.
Gophen, M. 1978. Errors in the estimation of recruitment of early stages of
Mesocyclops leuckarti (Claus) caused by the diurnal periodicity of egg-production.
Gophen, M., and R. Landau. 1977. Trophic interactions between zooplankton and
sardine Mirogrex terraesanctae populations in Lake Kinneret, Israel. Oikos 29:
Grogan, W.L., Jr., and L.J. Hribar. 2006. The bromeliad-inhabiting midge,
Forcipomyia (Phytohelea) bromelicola (Lutz), new to the fauna of the United
States. Entomological News 117:319–322.
Havel, J.E., C.E. Lee, and M.J. Vander Zanden. 2005. Do reservoirs facilitate invasions
into landscapes? BioScience 55:518–525.
Holyńska, M., J.W. Reid, and H. Ueda. 2003. Genus Mesocyclops Sars, 1914. Pp.
12–213, In H. Ueda, and J.W. Reid (Eds.). Copepoda: Cyclopoida. Genera
Mesocyclops and Thermocyclops. Guides to the Identification of the Microinvertebrates
of the Continental Waters of the World 20:1–318.
2008 L.J. Hribar and J.W. Reid 227
Ishida, T. 2002. Illustrated fauna of the freshwater cyclopoid copepods of Japan. The
Bulletin of the Biogeographical Society of Japan 57:37–106. [In Japanese, with
notes in English.]
Janicki, A., and J. DeCosta. 1990. An analysis of prey selection by Mesocyclops
edax. Hydrobiologia 198:133–139.
Karaytug, S., and G.A. Boxshall. 1998. Partial revision of Paracyclops Claus, 1893
(Copepoda, Cyclopoida, Cyclopidae) with descriptions of four new species. Bulletin
of the Natural History Museum, London, Zoology 64:111–205.
Krupa, E.G. 2005. First record of Mesocyclops ogunnus Onabamiro, 1957 from Kazakhstan
(Crustacea: Cyclopidae). Zoosystematica Rossica 14:23–26.
Lansac-Tôha, F.A., L.F.M. Velho, J. Higuti, and E.M. Takahashi. 2002. Cyclopidae
(Crustacea, Copepoda) from the upper Paraná River fl oodplain, Brazil. Brazilian
Journal of Biology 62:125–133.
Marten, G.G., and J.W. Reid. 2007. Cyclopoid copepods. In T.G. Floore (Ed.). Biorational
Control of Mosquitoes. American Mosquito Control Association Bulletin
7. Supplement to the Journal of the American Mosquito Control Association
Matsumura-Tundisi, T., and W.M. Silva. 2002. Occurrence of Mesocyclops ogunnus
Onabamiro, 1957 (Copepoda: Cyclopoida) in water bodies of São Paulo State,
identified as Mesocyclops kieferi Van de Velde, 1984. Brazilian Journal of Biology
Matsumura-Tundisi, T., A.C. Rietzler, E.L.G. Espíndola, J.G. Tundisi, and O. Rocha.
1990. Predation on Ceriodaphnia cornuta and Brachionus calycifl orus by two
Mesocyclops species coexisting in Barra Bonita reservoir (SP, Brazil). Hydrobiologia
Pagano, M., S. Kâ, and E.H. Ndour. 2003. Communautés zooplanctoniques:
Biomasses et taux de broutage. Pp. 59–73, In R. Arfi, N. Ba, M. Bouvy, C.
Corbin, Y. Diop, S. Kâ, F. Lebihan, M. Mboup, E.H. Ndour, M. Pagano, and S.
Sané. Lac de Guiers (Sénégal). Conditions Environnementales et Communautés
Planctoniques. Document Centre IRD, Dakar, Senegal. 77 pp.
Patalas, K. 1986. The geographical distribution of Mesocyclops edax (S.A. Forbes)
in lakes of Canada. Syllogeus 58:400–408.
Reid, J.W. 1999. New records of Bryocyclops from the continental USA, Puerto
Rico, and Brazil (Copepoda: Cyclopoida: Cyclopidae). Journal of Crustacean
Reid, J.W. 2000. Workshop on taxonomic techniques for copepods. “World of
Copepods” website. Department of Invertebrate Zoology, Smithsonian Museum
of Natural History. Available online at http://invertebrates.si.edu/copepod/
techniques.htm. Accessed 16 October 2007.
Reid, J.W. 2008. Arctodiaptomus dorsalis (Marsh): A case history of copepod dispersal.
Reid, J.W., and L.J. Hribar. 2006. Records of some Copepoda (Crustacea) from the
Florida Keys. Proceedings of the Academy of Natural Sciences of Philadelphia
Reid, J.W., and L. Moreno D. 1999. The western and southern distribution of Mesocyclops
edax. Proceedings of the Biological Society of Washington 112:581–591.
Reid, J.W., and R. Pinto-Coelho. 1994a. Planktonic Copepoda of Furnas Reservoir:
Initial survey of species (1993) and review of literature. Pp. 93–114, In R.M.
Pinto-Coelho, A. Giani, and E. von Sperling (Eds.). Ecology and Human Impact
on Lakes and Reservoirs in Minas Gerais with Special Reference to Future Development
and Management Strategies. SEGRAC, Belo Horizonte, Brazil.
228 Southeastern Naturalist Vol.7, No. 2
Reid, J.W., and R. Pinto-Coelho. 1994b. An Afro-Asian continental copepod, Mesocyclops
ogunnus, found in Brazil: With a new key to the species of Mesocyclops
in South America and a review of intercontinental introductions of copepods.
Reid, J.W., R. Hamilton IV, and R.M. Duffield. 2002. First confirmed New World
record of Apocyclops dengizicus (Lepeshkin), with a key to the species of Apocyclops
in North America and the Caribbean region (Crustacea: Copepoda: Cyclopidae).
Santos-Wisniewski, M.J., and O. Rocha. 2007. Spatial distribution and secondary
production of Copepoda in a tropical reservoir: Barra Bonita, SP, Brazil. Brazilian
Journal of Biology 67:223–233.
Service, M.W. 1993. Mosquito Ecology: Field Sampling Methods. Elsevier Applied
Science, London, UK. 988 pp.
Simberloff, D. 1997. The biology of invasions. Pp. 3–17, In D. Simberloff, D.C.
Schmitz, and T.C. Brown (Eds.). Strangers in Paradise: Impact and Management
of Nonindigenous Species in Florida. Island Press, Washington, DC. 467 pp.
Snyder, J.R., A. Herndon, and W.B. Robertson, Jr. 1990. South Florida Rockland. Pp.
230–277, In R.L. Myers and J.J. Ewel (Eds.). Ecosystems of Florida. University
of Central Florida Press, Orlando, FL. 765 pp.
Stern, W.L., and G.K. Brizicky. 1957. The woods and fl ora of the Florida Keys. Introduction.
Tropical Woods 107:36–65.
Suárez-Morales, E., J. McLelland, and J. Reid. 1999. The planktonic copepods of
coastal saline ponds of the Cayman Islands with special reference to the occurrence
of Mesocyclops ogunnus Onabamiro, an apparently introduced Afro-Asian
cyclopoid. Gulf Research Reports 11:51–55.
Tundisi, J.G., T. Matsumura-Tundisi, M.C. Calijuri, and E.M.L. Novo. 1991. Comparative
limnology of five reservoirs in the middle Tietê River, S. Paulo State.
Verhandlungen der Internationale Vereinigung für Theoretische und Angewandte
Warren, G.L. 1997. Nonindigenous freshwater invertebrates. Pp. 101–108, In D.
Simberloff, D.C Schmitz, and T.C. Brown (Eds.). Strangers in Paradise: Impact
and Management of Nonindigenous Species in Florida. Island Press, Washington,
DC. 467 pp.
Williamson, C.E. 1980. The predatory behavior of Mesocyclops edax: Predator
preferences, prey defenses, and starvation-induced changes. Limnology and
Williamson, C.E. 1984. Laboratory and field experiments on the feeding ecology of
the cyclopoid copepod, Mesocyclops edax. Freshwater Biology 14:575–585.
Williamson, C.E. 1986. The swimming and feeding behavior of Mesocyclops. Hydrobiologia
Wilson, M.S., and H.C. Yeatman. 1959. Free-living Copepoda. Pp. 735–868, In W.T.
Edmondson (Ed.). H.B. Ward and G.C. Whipple’s Freshwater Biology, 2nd Edition.
John Wiley and Sons, Inc., New York, NY. 1248 pp.
Yeatman, H.C. 1963. Some redescriptions and new records of littoral copepods for
the Woods Hole, Massachusetts region. Transactions of the American Microscopical
Zavortink, T.J., and G.F. O’Meara. 1999. Culex (Micraedes) biscaynensis n. sp. from
Florida (Diptera: Culicidae). Journal of the American Mosquito Control Association