William F. Loftus

HOW OFTEN DO FISHES “RUN ON EMPTY”?

We used a large data set of African, Neotropical, and North American fishes to examine the frequency with which fishes have empty stomachs (nspecies 5 254; nindividuals 5 36 875). Mean percentage of empty stomachs was low across all fishes (16.26 1.2%) but varied from 0% to 79.4% among individual species. Nocturnal fishes had empty stom- achs more frequently than diurnal fishes. Trophic classification was strongly associated with the percentage of empty stomachs, a pattern also revealed from an intraspecific analysis. Fishes appear to adjust their feeding intervals relative to the energy density, conversion efficiency, and particle size of their food. Piscivorous fishes seem to be the only trophic group that regularly experience long periods of empty stomachs, with species that consume prey whole and those that provide extended parental care having the highest proportions of empty stomachs. Activity patterns and life histories of some piscivorous species probably have evolved in partial response to energetic benefits of large, energy-rich food resources.

Empirical assessment of fish introductions in a subtropical wetland: An evaluation of contrasting views

We summarized data from eight quantitative fish surveys conducted in southern Florida to evaluate the distribution and relative abundance of introduced fishes across a variety of habitats. These surveys encompassed marsh and canal habitats throughout most of the Everglades region, including the mangrove fringe of Florida Bay. Two studies provided systematically collected density information over a 20-year period, and documented the first local appearance of four introduced fishes based on their repeated absence in prior surveys. Those species displayed a pattern of rapid population growth followed by decline, then persistence at lower densities. Estuarine areas in the southern Everglades, characterized by natural tidal creeks surrounded by mangrove-dominated marshes, and canals held the largest introduced-fish populations. Introduced fishes were also common, at times exceeding 50% of the fish community, in solution holes that serve as dry-season refuges in short-hydroperiod rockland habitats of the eastern Everglades. Wet prairies and alligator ponds distant from canals generally held few individuals of introduced fishes. These patterns suggest that the introduced fishes in southern Florida at present may not be well-adapted to persist in freshwater marshes of the Everglades, possibly because of an interaction of periodic cold-temperature stress and hydrologic fluctuation. Our analyses indicated low densities of these fishes in central or northern Everglades wet-prairie communities, and, in the absence of experimental data, little evidence of biotic effects in this spatially extensive habitat. There is no guarantee that this condition will be maintained, especially under the cumulative effects of future invasions or environmental change.

Long-term hydrologic effects on marsh plant community structure in the southern Everglades

Although large-scale transformation of Everglades landscapes has occurred during the past century, the patterns of association among hydrologic factors and southern Everglades freshwater marsh vegetation have not been well-defined. We used a 10-year data base on the aquatic biota of Shark Slough to classify vegetation and describe plant community change in intermediate- to long-hydroperiod Everglades marshes. Study area marsh vegetation was quantitatively grouped into associations dominated by 1)Cladium jamaicense, 2) a group of emergents includingEleocharis cellulosa, Sagittaria lancifolia, andRhyncospora tracyi, 3) taxa associated with algal mats (Utricularia spp. andBacopa caroliniana), and 4) the grassesPanicum hemitomon andPaspalidium geminatum. During the decade evaluated, the range of water depths that characterized our study sites approached both extremes depicted in the 40-year hydrologic record for the region. Water depths were near the long-term average during the mid-1980s, declined sharply during a late 1980s drought, and underwent a prolonged increase from 1991 through 1995. Overall macrophyte cover varied inversely with water depth, while the response of periphyton was more complex. An ordination analysis, based on plant species abundance, revealed that study area vegetation structure was associated with hydrologic patterns. Marsh plant community structure showed evidence of cyclic interannual variation corresponding to hydrologic change over the decade evaluated. Lower water depths, the occurrence of marl substrates, and high periphyton cover were correlated. These factors contributed to reduced macrophyte cover in portions of the study area from which water had been diverted.

Community structure of fishes inhabiting aquatic refuges in a threatened Karst wetland and its implications for ecosystem management

Abstract We illustrate the importance of subsurface refuges for conservation of aquatic fauna with our studies of karstic wetlands in Everglades National Park, Florida, USA. Managers have proposed that water levels there should not fall more than 46 cm below ground level for more than 90 days annually. In four areas, 84% of solution holes were less than 46 cm deep and holes deeper than 1m were rare (

SOUTHERN MARL PRAIRIES CONCEPTUAL ECOLOGICAL MODEL

About 190,000 ha of higher-elevation marl prairies flank either side of Shark River Slough in the southern Everglades. Water levels typically drop below the ground surface each year in this landscape. Consequently, peat soil accretion is inhibited, and substrates consist either of calcitic marl produced by algal periphyton mats or exposed limestone bedrock. The southern marl prairies support complex mosaics of wet prairie, sawgrass sawgrass (Cladium jamaicense), tree islands, and tropical hammock communities and a high diversity of plant species. However, relatively short hydroperiods and annual dry downs provide stressful conditions for aquatic fauna, affecting survival in the dry season when surface water is absent. Here, we present a conceptual ecological model developed for this landscape through scientific concensus, use of empirical data, and modeling. The two major societal drivers affecting the southern marl prairies are water management practices and agricultural and urban development. These drivers lead to five groups of ecosystem stressors: loss of spatial extent and connectivity, shortened hydroperiod and increased drought severity, extended hydroperiod and drying pattern reversals, introduction and spread of non-native trees, and introduction and spread of non-native fishes. Major ecological attributes include periphyton mats, plant species diversity and community mosaic, Cape Sable seaside sparrow (Ammodramus maritimus mirabilis), marsh fishes and associated aquatic fauna prey base, American alligator (Alligator mississippiensis), and wading bird early dry season foraging. Water management and development are hypothesized to have a negative effect on the ecological attributes of the southern marl prairies in the following ways. Periphyton mats have decreased in cover in areas where hydroperiod has been significantly reduced and changed in community composition due to inverse responses to increased nutrient availability. Plant species diversity and community mosaics have changed due to shifting gradients to more terrestrial or more aquatic communities, displacement of native communities by non-natives, expansion of woody plants, high-intensity dry season fires, tree-island burnout, and reduced microtopography resulting from alligator population decline. Cape Sable seaside sparrow populations are threatened by nest destruction resulting from extended hydroperiods, drying pattern reversals, and high intensity dry season fires, as well as by the expansion of woody plants into graminoid wetland habitats. Populations of marsh fishes and associated aquatic fauna that constitute the aquatic prey base for higher vertebrates have decreased in density and changed in species composition and size structure due to loss of wetland spatial extent, shortened hydroperiod, increased drought severity, loss of aquatic drought refugia in solution holes and alligator holes, and spread of exotic fishes. American alligator populations have declined in the Rocky Glades, and alligator holes have filled with sediment, as a result of shortened hydroperiod and increased drought severity. Habitat options for wading birds to forage during the early dry season and during unusually wet years have been reduced due to loss of spatial extent and shortened hydroperiod.

Modeling fish dynamics and effects of stress in a hydrologically pulsed ecosystem

Many wetlands undergo seasonal cycles in precipitation and water depth.This environmental seasonality is echoed in patterns of production of fishbiomass, which, in turn, influence the phenology of other components of thefood web, including wading birds. Human activities, such as drainage orother alterations of the hydrology, can exacerbate these natural cycles andresult in detrimental stresses on fish production and the higher trophic levels dependent on this production. In this paper we model theseasonal pattern of fish production in a freshwater marsh, with specialreference to the Everglades/Big Cypress region of southern Florida.The model illustrates the temporal pattern of production through theyear, which can result in very high densities of fish at the end of ahydroperiod (period of flooding), aswell as the importance of ponds and other deep depressions, both as refugia and sinks during dry periods. The model predicts that: (1) there is an effective threshold in the length of the hydroperiod that must beexceeded for high fish-population densities to be produced, (2) large,piscivorous fishes do not appear tohave a major impact on smaller fishes in the marsh habitat, and (3) therecovery of small-fish populations in the marsh following a major droughtmay require up to a year. The last of these results is relevant toassessing anthropogenic impacts on marsh production, as these effectsmay increase the severity and frequency of droughts.

Distribution and relative abundance of the crayfishes Procambarus alleni (Faxon) and P. fallax (Hagen) in southern Florida

The Everglades crayfish (Procambarus alleni Faxon) is the only species reported from Everglades National Park (ENP) and Big Cypress National Preserve (BICY). However, we identified a second species, Procambarus fallax (Hagen), from those locations. Reexamination of archived samples showed that P. fallax had been collected as early as 1985 but had been misidentified or overlooked; therefore, we provide methods for distinguishing live and preserved specimens of these morphologically similar species. Species relative abundance varied predictably with hydroperiod (length of inundation) both in recent collections and in archived samples. Procambarus fallax relative abundance was highest in sites characterized by long hydroperiod and prolonged inundation, whereas P. alleni was the lone species in short hydroperiod marshes.

Diapause in copepods (Crustacea) from ephemeral habitats with different hydroperiods in Everglades National Park (Florida, U.S.A.)

Water management practices in the Everglades have severely stressed the natural system, particularly by reducing the hydroperiods of much of the region. During the dry season of 1999, we investigated the influence of hydroperiod on the species composition and dormancy patterns of freshwater copepod communities in seasonal wetlands of Everglades National Park, Florida, U.S.A. The habitats were characterized by an annual dry season, from December through June. We sampled at two locations: the Long Pine Key area of the Rocky Glades region (short hydroperiod, ca. 4–5 months), and western Taylor Slough (intermediate hydroperiod, ca. 8–10 months). Both areas have experienced a reduction in natural hydroperiods and an increase in the frequency of dry-down. We collected weekly plankton samples from Rocky Glades solution holes to assess the potential species pool of copepods. To document the taxa capable of surviving dry-down by resting, we performed three immersion trials in which we rehydrated, in laboratory aquaria, sediment patches from solution holes and surface soils from all stations. Only a subset of the planktonic species collected emerged from the dried sediments. The cyclopoids Microcyclops rubellus and Paracyclops poppei were dominant. This is the first record of diapause for P. poppei. Species distributions from the different hydroperiod soil patches indicated that more diapausing species occurred at the sites that dried for shorter periods. Emerging individuals of M. rubellus and P. poppei were mainly ovigerous females, demonstrating a resting strategy seldom before recorded. The cyclopoid Diacyclops nearcticus had not been previously reported to diapause, but they emerged from the dried sediments in our trials. Our collections included six new records for Florida: Diacyclops nearcticus, Megacyclops latipes, Orthocyclops modestus, Elaphoidella marjoryae, Bryocamptus sp. and Bryocamptus cf. newyorkensis. Paracyclops poppei, Macrocyclops fuscus and Arctodiaptomus floridanus are new records for Everglades National Park. Clearly, diapause is an important strategy for the persistence of copepods in short-hydroperiod wetlands. The duration of the dry period appears to be inversely related to the number of species that emerge from diapause.

Salinity Tolerance of the African Jewelfish Hemichromis letourneuxi, a Non-native Cichlid in South Florida (USA)

Abstract The African Jewelfish (Cichlidae: Hemichromis letourneuxi) is a predatory, non-native fish that has recently (since 2000) begun to expand its geographic range across south Florida. The salinity tolerance of H. letourneuxi was unknown, and thus it was unclear whether the species could use estuarine or coastal environments. The response of H. letourneuxi to chronic change in salinity was evaluated here by exposing fish to progressively increasing salinities (0–80 ppt). Fish were held at target salinities for a minimum of 30 days. The species showed excellent survival from 0–50 ppt. At 60 ppt, only 25% of the fish survived, and mean estimated survival time was 12 days. Above 60 ppt, mortality was 100%. Fish grew equally well from 0–50 ppt. In another experiment, fish were transferred directly from freshwater to various salinities from 5–35 ppt (seawater) and held for seven days, after which survivors were returned to freshwater. All fish transferred directly from freshwater to salinities up to 20 ppt survived; only 56% survived when transferred from freshwater to 25 ppt, and none survived transfer above 25 ppt. Experimental results indicated that H. letourneuxi can persist easily in salinities prevalent in coastal environments, even during periods of hypersalinity common in south Florida estuaries. Salinity will not restrict its dispersal by coastal pathways.

Recent Fish Introductions Into Everglades National Park: An Unforeseen Consequence of Water Management?

Non-native fishes present a management challenge to maintaining Everglades National Park (ENP) in a natural state. We summarized data from long-term fish monitoring studies in ENP and reviewed the timing of introductions relative to water-management changes. Beginning in the early 1950s, management actions have added canals, altered wetland habitats by flooding and drainage, and changed inflows into ENP, particularly in the Taylor Slough/C-111 basin and Rocky Glades. The first non-native fishes likely entered ENP by the late 1960s, but species numbers increased sharply in the early 1980s when new water-management actions were implemented. After 1999, eight non-native species and three native species, all previously recorded outside of Park boundaries, were found for the first time in ENP. Several of these incursions occurred following structural and operational changes that redirected water deliveries to wetlands open to the eastern boundary canals. Once established, control non-native fishes in Everglades wetlands is difficult; therefore, preventing introductions is key to their management. Integrating actions that minimize the spread of non-native species into protected natural areas into the adaptive management process for planning, development, and operation of water-management features may help to achieve the full suite of objectives for Everglades restoration.