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Temporal and Spatial Variations inCopepod (Crustacea) Communities inGroundwater in the Rocky Glades ofEverglades National Park (Florida, USA)M. Cristina Bruno a & Sue A. Perry aa South Florida Natural Resources Center, Everglades Nationsl Park,40001 State Road 9336, Homestead, Florida, 33034, USA E-mail:Published online: 11 Jan 2011.

To cite this article: M. Cristina Bruno & Sue A. Perry (2005) Temporal and Spatial Variations inCopepod (Crustacea) Communities in Groundwater in the Rocky Glades of Everglades National Park(Florida, USA), Journal of Freshwater Ecology, 20:1, 27-36, DOI: 10.1080/02705060.2005.9664933

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Temporal and Spatial Variations in Copepod (Crustacea) Communities in Groundwater in the Rocky Glades of

Everglades National Park (Florida, USA) M. Cristina Bruno and Sue A. Perry

South F/onUa Natuml Resou~es Cent84 Everg/ades Nations/ Pafit 40001 State Rasd9336, Homesteai.? F/orida 33034 USA

ABSTRACT We studied species composition and individual abundance of copepods in the

surficial aquifer in short-hydroperiod habitats of Everglades National Park by collecting copepods from different depths in wells for three consecutive years. The wells were cased and open at depths that corresponded to highly permeable layers. Groundwater communities were dominated by surface copepods which colonized groundwater mainly during the dry season. The total number of copepods collected decreased exponentially with depth; the decrease in copepod numbers and species richness below the 3 m depth was due to high permeability of the limestone above 3 m depth and to the presence of a semipermeable layer at lower depths. The calanoid Osphranticum labronecrum and the cyclopoids Orthocyclops modestus and Thennocyclops parvus that were dominant in the collections can be considered stygophiles at least in Everglades National Park. Copepod groundwater communities were most similar on a local scale, indicating that when local surface water populations enter groundwater by following the receding water table, they do not disperse widely through the groundwater system. Densities of groundwater populations of stygophiles were low, which increases their risk of being impacted by changes in hydrology.

INTRODUCTION In Everglades National Park (ENP), marsh hydrology changed radically during the

1950s and 1960s as a result of the Central and South Florida Project, which impounded and diverted water from the north, draining it eastward and southward by canals. Water depths and hydroperiods in ENP were decreased dramatically (Loftus et al. 1992). Particularly affected by drainage and insufficient water deliveries were the higher elevation Rockland marshes, or Rocky Glades, a once-large karstic system that is now confined within ENP between Shark River and Taylor sloughs. Reduced hydroperiods and lower groundwater levels in the Rocky Glades have greatly diminished aquatic productivity and the capacity of its karstic features to serve as a refugium for aquatic life during the dry season (Loftus et al. 1992).

In the Rocky Glades, a high degree of dissolution of the oolitic limestone bedrock has occurred with time, producing a typical karstic landscape with thousands of solution holes (Hoffmeister 1974), which provide a vertical dimension of habitat for aquatic organisms. During the wet season (June-October), rainfall and groundwater-recharge fill the solution holes, and lower-elevation areas of the Rocky Glades can be flooded in years of high rainfall. In the dry season (November-May), surface water disappears as a result of evaporation, percolation, and evapotranspiration.

In the southern Everglades, limestone of the Miami Limestone and Fort Thompson Formation form the Biscayne aquifer in the upper part of the surficial aquifer. In ENP, the Fort Thompson Formation is 3-15 m deep, and it thickens slightly to the east, where it underlies the Miami Limestone (Fish and Stewart 1991). The Miami Limestone and Fort Thompson Formation consist of five time-stratigraphic marine units termed, from oldest to youngest, Q1 through Q5 (Q for Quaternary; Perkins 1977). Earlier investigators presented evidence for at least one unit of very limited ground water flow within the Biscayne Aquifer

Journal of Freshwater Ecology, Volume 20, Number 1 - March 2005

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(Cunningham and Wright 1998, Genereux and Guardiario 1998, Kaufman and Switanek 1998, Nemeth et al. 2000, Sonenshein 2001). at the top of the Q3 in the Fort Thompson Formation (Genereux and Guardiario 1998).

Over the past five years, we have been conducting extensive surveys of copepod communities from surface water in ENP (Bruno et al. 2001 and 2002b) and from groundwater in areas of Dade County east of the park (Bruno et al. 2003a) to document the fauna and to better understand the dynamics and relationships with hydrological regimes. Groundwater copepod communities in the Rocky Glades were also studied (Bruno and Perry 2004) from wells where a continuous exchange between surface and groundwater fauna was possible. In this study, we examined the composition and dynamics of groundwater communities in the Rocky Glades over a three-year time period from wells where the faunal exchange between surface and groundwater was reduced by well features. Our objectives were: 1) to define the stygoxenic and stygophilic copepod species; 2) to assess how seasonal changes in groundwater levels influence the ability of copepods to disperse into groundwater; and, 3) to verify how local geological features influence faunal exchange between surface and groundwater.

MATERIALS AND METHODS

Two sites (site 1 : 25"23'59.7"N, 080°38'23.4"W, site 2: 25'25'21 .8"N, 080°42' 15.5"W) were selected in the Rocky Glades in areas with high porosity identified by Cunningham (2004) in spring 2000 with a scanning penetrating radar. Three wells were cored (CHI, CH2, CH3). and a hydrogeological study of the area was conducted by Cunningham (2004). On the basis of these results, three additional monitoring wells (MW) were drilled, and cased from the surface to those high porosity zones (Table 1). Quantitative samples were collected from the wells using a portable pump and PVC pipes extended to a depth within or immediately above the high porosity layer. At MW4, the suction pipe extended only to 3 m because of a partial obstruction of this well. One thousand liters of water were filtered using a 40-pm mesh plankton net. Water depth was measured with a meter tape. Temperature, conductivity, and oxygen percent saturation were recorded using a YSI 85@ meter. Rainfall was recorded at ENP monitoring stations RPLl and RPL2 (25"23'04.78"N, 80°35'38.01"W). Because rainfall is the main recharge source for the surficial aquifer in the Rocky Glades, when analyzing the relationship between monthly groundwater levels and rainfall, we used the total monthly rainfall between the sampling date and the sampling date of the previous month.

Table 1. Characteristics of the wells, and of the wells area, and sampling depths for each well.

Site Well Well Casing Open hole High porosity Sampling name depth (m) depth (m) (m) layer depth (m) depth (m)

1 MW5 3 2.25 2.25-3 2.5 2.5 m 1 MW4 4.5 3.045 3.045-4.5 4 2.5 m 1 MW9 11.10 5.25 5.25-11.10 6,7.5,9 5.5.7, 8.5 m 1 CH 1 11.55 6.3 6.3-1 1.55 7-8 8.1 m 1 CH2 12.70 2.25 2.25-12.70 7-9, 12 8.1, 11.1 m 2 MW6 3 2.25 2.25-3 3.5 2.5 m 2 MW7 4.875 3 3-4.875 3.5 2.5 m 2 MW8 6.27 5.13 5.13-6.27 5.5 5.5 m 2 CH3 9.6 6.6 6.6-9.6 7-8 7

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Twelve samples were collected monthly, at the beginning of each month, from May 2001 to April 2004. One sampling year started when groundwater levels were at their lowest point at the end of the dry season (May), to begin the sampling year at the start of the wet season. Sampling years were thus defined as first year(May 2001 to April 2002), second year (May 2002 to April 2003). and third year (May 2003 to April 2004).

During four weeks in July 2001, we set one micro-trap baited with shrimp at each well and retrieved it after 24 hours. This trapping was repeated for four weeks in October 2001. Traps were ineffective, collecting only two copepods, both in July. A male of Paramphiascella sp. was collected from MW8 at 6 m depth, and a female of Nitokra biserosa Mielke 1993 was collected from MW5 at 3 rn depth. These faunal results were not included in the quantitative analysis.

All the faunal samples were fixed in the field with 5% buffered formalin. Specimens were sorted and counted under magnification, mounted on permanent slides, and examined with phase contrast microscopy. All copepod identifications included sex and developmental stage and followed Yeatman (1959). Dussart (1967), SuQez-Morales et al. (1996), Reid (1992), Karaytug (1999), Bruno et al. (2000,2002a). Adult and copepodites stage IV and V were identified to the species level. Individuals of earlier copepodite stages and all nauplii were not identified, but were collectively labelled "copepodites" and "nauplii". Nauplii were not used in the statistical analyses since they could have been overlooked, given their small sizes.

We ran multiple correlations among groundwater depth, temperature, oxygen, conductivity, number of individuals, and number of species collected at each well each month from May 2001 to April 2004. We also ran multiple correlations among the average values of groundwater depth, temperature, oxygen, conductivity, total rainfall, the total number of individuals and the total number of species recorded monthly from May 2001 to April 2004. We analyzed the spatial distribution of copepods with a detrended correspondence analysis of the total number of copepods of each taxon collected monthly at each well, transformed with Log (x+l).

RESULTS The lowest groundwater levels were reached at the end of the dry season in May for

2001(-98 cm) and 2002 (-1 39 cm) and in April for 2004 (-1 15 cm) (Fig. 1). The highest water levels were reached in October 2001 (-13 cm), July 2002 (-15 cm), and October 2003 (-9 cm), within two months following the peak in summer rainfall. As a consequence of the different rainfall patterns for the three sampling years, groundwater levels were high only from August to December in 2001, from July to September in 2002, and from June to November in 2003. Total monthly rainfall between sampling dates and average groundwater levels at the time of sampling were strongly correlated ( ~ 0 . 5 5 , p<0.01). Groundwater depth was correlated with conductivity (r=0.10, p=0.033) and temperature (r=-0.34, p<0.001).

Conductivity ranged from 204 pS in April 2003 to 1271 pS. Average monthly values were correlated with total monthly rainfall (p=-0.33. p=0.047). Monthly temperature ranged between 22.3 "C and 26.5 "C and decreased with increasing depth. Dissolved oxygen was always low (average: 1.4 % saturation) and ranged from <0.1 to 35% saturation. Oxygen percent saturation decreased with increasing depth.

We collected a total of 2,18 1 copepods (Table 2), belonging to 14 species, seven of which were rare (less than 10 specimens collected over the entire sampling period). Most of the species had been previously collected in groundwater in ENP (Bruno and Perry 2004). The number of individuals and number of species collected each month for each sample were significantly correlated (r=0.47, p<0.001); the number of species was also correlated with conductivity (r=-0.11, p=0.025) and with temperature (r=0.29, p<0.001). In addition,

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the monthly average number of individuals and number of species were correlated with average monthly groundwater levels (p0 .35 , p=0.036, and I=-0.33, p=0.048, respectively).

Higher numbers of copepods were collected at site 2 than at site 1 (Fig. 2). Total numbers of copepods collected at each depth (3,6,7.5,9, and 12 m) were significantly different (Kruskall-Wallis, peO.00) and decreased exponentially with depth (Fig. 3). The number of copepods collected at the 3 and 7.5 m depth was significantly different from the number of copepods collected at all the other depths (Mann-Whitney, p<0.05).

The number of individuals collected each month varied among the three years (418 in 2001, 1404 in 2002, and 359 in 2003) and seasonally within each year. The highest total number of individuals was collected in sampling year 2002 (65% of the three years total), followed by 2001 (19%) and 2003 (16%). Years 2001 and 2003 had similar range and variation in number of individuals; sampling year 2002 had higher numbers during the dry season and lower numbers in the wet season than the other two sampling years. Each year, numbers were higher during the dry season. Species richness was higher the first sampling year, and decreased with time, mostly because rare species disappeared. The rare species

-Total rainfall (L) +Average groundwater level (R)

Figure 1. Average groundwater level at sampling date and total rainfall recorded between the sampling date and the previous one

Table 2. Total numbers of individuals collected for each taxon.

Taxon Total Collected Ecological Ecological number of elsewhere in classification in ENP classification in ENP individuals ENP in (after Bruno et al. (present study)

Orthocyclops modestus l o 0 0 Diacyclops nearcticus 419 Thennocyclops parvus 313 Osphranricum kabronectum 129 Copepodites 107 Nauplii 82 Microcyclops rubellus 71 Macrocyclops albidus 30 Microcyclops vancans 15 Acanrhocyclops robustus 4 Phyllognothopus viguien 4 Mesocyclops amencanus 2 Tropocyclops prasinus mexicanus 2 Paracyclops poppei 1 Onychohnmptus mohammed 1

groundwater Yes Yes Yes Yes Yes Yes Yes Yes No Yes Yes Yes Yes No No

2004) Epigean, stygoxene Stygophile Epigean, stygoxene Epigean, stygoxene n.a. n.a. Epigean. stygoxene Epigean. stygoxene

Epigean, stygoxene Epigean. stygoxene Epigean. stygoxene Epigean. stygoxene

Stygophile Stygophile Stygophile Stygophile n.a. n.a. Epigean, stygoxene Epigean, stygoxene Epigean, stygoxene Epigean. stygoxene Epigean. stygoxene Epigean. stygoxene Epigean. stygoxene Epigean. stygoxene Epigean, stygoxene

Ameiridae 1 No unknown

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were collected mostly during the first year, and again in October 2002 and March 2003. From July 2003 on, 92% of all the individuals collected belonged to three dominant species - Themcyclops parvus Reid 1989, Diacyclops nearcticus Kiefer 1934, and Orthocyclops modestus (Herrick 1883).

The species composition (Table 2) was similar to that recorded for a different set of wells in the Rocky Glades by Bruno and Perry (2004); the first four ranked species of this study (0. modestus, D. nearcticus, T. parvus, and Osphranticum labronectum S.A. Forbes 1882) were dominant in the other Rocky Glades wells previously studied. In this study, 0. modestus was present mostly in the dry season, and T. parvus and D. nearcticus mostly in the wet season. Peaks in number of individuals for all three species corresponded to high rainfall months. All the species recorded are found in surface water in ENP and elsewhere.

The first three axes for the detrended correspondence analysis had total eigenvalue of 0.79. On the biplot of the first two axes (Fig. 4). the wells with the highest number of individuals collected, all belonging to the abundant species, were grouped near the center of the graph, whereas those wells with few specimens were separated at the extremities of both axes. At well MW9 we collected few common species and three of the five very rare ones Onychocamptus mohammed (Blanchard and Richard 1891), Paracyclops poppei (Rehberg 1880), Tropocyclops praxinus mexicanus Kiefer 1938. The other two rare species were collected at MW5 (one unidentified copepodite, family Ameridae) and at CHI and MW7 (Mesocyclops amencanus Dussart 1985, two individuals). Axis 3 (Fig. 5) records similar species composition for wells located near each other; wells were separated on the basis of the geographical position as wells at site 1 (CHI, CH2, MW4, MW5, MW9), and at site 2 (CH3, MW6, MW7, MW8). The common and abundant species were grouped with site 2 wells, whereas the rare species were grouped with site 1 wells.

Figure 2. Average number of copepods collected at each well. Black: site 1; white: site 2.

DISCUSSION According to Cunningham (2004), who analyzed the porosity and hydraulic

conductivity at CH2 and MW9, the Q5 (0-2 m below surface) has very high porosity and hydraulic conductivity, whereas the Q4 (from 2 to 4 m below surface) has lower porosity and hydraulic conductivity. As a consequence, the freshwater content is higher in the Q5 than in the Q4. However, both the Q4 and the Q3 (from 4 to 7.5 m below ground) have higher porosity near the bottom of the layer. The decrease in copepod numbers and species richness below the Q4 recorded in this study might be due to the high permeability in Q5, which could act as a preferential passageway for epigean forms, and to the presence of the

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semipermeable layer in 4 4 and upper 43, which could act to retard vertical organism migration. Copepods were most abundant at the 3 and 7.5 m depths, in the Q5 and bottom of the Q4 and 43. The study of the coreholes and optical images collected at site 2, where most of the copepods were collected, was not completed (Cunningham, pen. corn.). The two wells where high numbers of copepods were collected, reached either the Q5 unit (MW6), and the Q4 (MW7). These layers are generally well-connected to surface water. The connections to surface water explain the high faunal exchange between surface and ground water and thus the high number of copepods collected at those wells. Why the shallow wells at site 1, which also reach the Q5 unit, had low counts is uncertain but could be due to vertical movement of deeper ground water toward the surface, as imaged with the ground penetrating radar along the road adjacent to Site 1 (Cunningham, pers. com.). If this occurs, deeper groundwater. typically poor in fauna, could dilute the shallow Q5 ground water, which is well connected to surface water and richer in fauna. Although we collected high numbers of individuals at site 2, probably due to the higher permeability of the limestone, groundwater copepod communities at site 1 were more diverse.

Copepods appear to have the potential ability to disperse passively over long distances when they follow the movement of groundwater, especially in areas that connect with lower elevation and therefore have faster groundwater flow. Two specimens collected in summer 2001 with baited traps are associated with marine habitats. These organisms were possibly transported into groundwater in the Rocky Glades by saltwater intrusion. Saltwater intrusion in the shallow aquifer has been reported for the entire surficial aquifer in ENP (Price and Swart 2000). These faunal findings, together with the high values of conductivity we recorded in a well at site 2 are consistent with saltwater intrusion and mixing with surface water at low depths in the aquifer.

In a previous study (Bruno and Perry 2004) on another set of wells in the Rocky Glades, the temporal distribution of the species that were most abundant was related to seasonal abundance in surface waters, which supports the hypothesis of colonization by surface taxa into groundwater throughout the year. Colonization of the surficial aquifer in the Rocky Glades by surface copepod taxa peaked at the beginning of the wet season when flooding passively transported surface water copepods to new areas. Colonization by surface taxa also occurred at the end of the wet season when they were passively or actively transported into groundwater by the receding water table. In the same study, when groundwater levels dropped about 60 cm below ground level, the density of individuals and species richness dropped, suggesting that the ability of copepods to migrate vertically into the aquifer or to survive adverse physical and chemical conditions in groundwater was

Figure 3. Average number of copepods collected at each depth. Line represents nonlinear regression (exponential equation).

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reduced at low groundwater levels. Those results are in contrast to what we report here, where there was an inverse correlation between groundwater levels and number of individuals and of species, and where numbers tended to be higher during the dry season. Copepod average density in this study was 104 times lower than the average density of the previous study (Bruno and Perry. 2004). The important difference between the two studies is the structure of the wells. In the first study, the wells were deep and not cased, therefore vertical and horizontal movements of copepods were possible. Copepods were able to migrate both vertically in the well water column or through the limestone vertically and horizontally into the well. In the present study, because of the casing, only copepods that were present in the aquifer at the sampled depth were collected. Of the 23 species reported by Bruno and Perry. (2004) for groundwater in the Rocky Glades, 10 were collected in the present study, and the four rare species in this study were not previously collected. All of the most abundant species of copepods collected in groundwater by Bruno and Perry. (2004) were described as stygoxenes, few true stygophile and stygobite taxa were present, and most of them were rare. The results of this study give further information on the ability of the surface taxa to colonize groundwater in the Rocky Glades. Some taxa such as the calanoid ArctodiaptomusJIoridanus (Marsh 1926) and the cyclopoid Eucyclops conrowae Reid 1992 are probably passively transported into the groundwater system when groundwater recedes at the end of the wet season. They are true stygoxenes (i.e., organisms with no affinities for the groundwater system but which occur accidentally in it). In fact, these taxa were abundant in the uncapped wells in the Rocky Glades but were not collected in the present study. The only other calanoid reported for ENP (Bruno et al. 2003a). 0. labronectum, and the cyclopoids 0. modestus and T. parwus were dominant in this study collection. These taxa are displaced from surface water, and transported into groundwater during high rainfall events, where they can survive even with reduced exchange with surface water. These species can be considered stygophiies (i.e., organisms that have greater affinities with the groundwater environment than the stygoxenes). They actively exploit resources in the groundwater system, and actively seek protection from unfavorable situations in the surface environment resulting from biotic or abiotic processes (Gibert et al. 1994), at least in ENP.

-.

A M . albidus

~M.amcricanus

MWB-6 CHI-9

I

Figure 4. Delrended correspondence analysis, ordination byplot for axes 1 and 2. Well locations are identified as MW- and CH-; the second number listed indicates the sampling depth for each well.

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Copepods were collected mainly at the surface/groundwater ecotone. A sharp decline in species richness of invertebrate communities with increasing depth into groundwater is a common phenomenon (see review in Strayer 1994). Several environmental factors control the distribution of groundwater biota, the most important one is probably the oxygen concentration (Danielopol and Niederreiter 1987). Surface water and groundwater interface sites have intense hydraulic exchanges. The biogeochemical activity at these interfaces is higher than in neighboring surface and groundwater systems (Gibert et al. 1997). and provides more trophic resources for copepods. However, hydraulic conductivity determines renewal rates of water and dissolved substance (such as oxygen, and organic carbon) to an aquifer (Strayer 1994) and also reflects the degree of connections of pores that serve as physical habitats for groundwater organisms. The different permeability of limestone layers reported by Cunningham (2004) is probably affecting the local functioning of the entire subsurface habitat.

Groundwater copepod communities were more similar on a local scale. When local surface water populations enter the shallow aquifer by following the receding water table, they do not generally disperse widely through the groundwater system. The dissimilarity in communities over larger distances may reflect differences in surface habitats as well as limited dispersal due to different porosity of the limestone. Previous studies in the limestone of the Rocky Glades (Bruno and Perry. 2004) and in adjacent areas (Bruno et al. 2003a) showed low dispersal for local groundwater populations. Densities of groundwater populations of stygophiles in ENP are low, and the copepod populations are at risk of being impacted by changes in hydrology. Lower groundwater levels during the dry season or reduced surface inundation during the wet season could result in reduced dispersal, increased populations isolation, and habitat fragmentation, which would increase the risk of extinction of local populations.

Figure 5. Detrended correspondence analysis, ordination byplot for axes 2 and 3. Well locations are identified as MW- and CH-; the second number listed indicates the sampling depth for each well.

ACKNOWLEDGEMENTS We thank William Loftus (USGSIBRD) for financing the wells drilling, Kevin

Cunningham (USGSIWRD) for providing information on the geological settings of the study area. The ENP volunteers Samuel Alvarenga, Carolina Ansani, Alessandro

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Campanaro, Veronica Canales, Anne Marie Labouche, Lorena Morales, Diogo Moreira, Roberta Raschioni, Timothy Schmidt, Fernanda de Sousa, and Florian Veau helped M.C. Bruno with fieldwork. This research was supported by South Florida Critical Ecosystem Science Initiative funding to the PI, S.A. Perry. Funding for M.C. Bruno was made possible by cooperative agreement no 5280-8-9015 with Kelsey Downum, Florida International University, Miami, Florida.

LITERATURE CITED Bruno, M. C. and S. A. Perry. 2004. Exchanges of copepod fauna between surface- and

groundwater in the Rocky Glades of Everglades National Park (Florida, U.S.A.). Arch Hydrobiol. 159(4): 489-510.

Bruno, M. C, J. W. Reid, and S. A. Perry. 2000. New records of copepods from Everglades National Park (Florida, U.S.A), descriptions of Elaphoidella marjoryae sp. nov., of ElaphoidellaJluviusherbae sp. nov. (Harpacticoida, Canthocamptidae), and supplementary description of Diacyclops nearcticus (Kiefer, 1934) (Cyclopoida, Cyclopidae). Crustaceans 73 (10): 1 17 1- 1205.

Bruno, M. C., W. F. Loftus, J. W. Reid, and S. A. Perry. 2001. Diapause in copepods (Crustacea) from ephemeral habitats with different hydroperiods in Everglades National Park (Florida, USA), p. 295-308 In: R. M. Lopez, J. W. Reid, and C. E. F. Rocha (eds.). Copepoda: Developments in ecology, biology and systematics. Proceedings of the 7' International Conference on Copepods, Curitiba, Brazil, 24-31 August 1999. Dev. Hydrobiol. 453/454.

Bruno, M. C, J . W. Reid, and S. A. Perry. 2002a. New records of copepods from freshwater in Everglades National Park (Florida): description of Nitokra evergladensis sp. nov. (Harpacticoida, Ameiridae), redescription of Bryocamptus newyorkensis (Chappuis 1926) (Harpacticoida, Canthocamptidae), and supplementary description of Attheyella americana (Herrick 1884). J . Crustacean Biol. 22 (4):834-854.

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Received- 30 April 2004 Accepted: 18 August 2004

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