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.     

Spatial patterns of fish communities along two estuarine gradients in southern Florida

In tropical and subtropical estuaries, gradients of primary productivity and salinity are generally invoked to explain patterns in community structure and standing crops of fishes. We documented spatial and temporal patterns in fish community structure and standing crops along salinity and nutrient gradients in two subtropical drainages of Everglades National Park, USA. The Shark River drains into the Gulf of Mexico and experiences diurnal tides carrying relatively nutrient enriched waters, while Taylor River is more hydrologically isolated by the oligohaline Florida Bay and experiences no discernable lunar tides. We hypothesized that the more nutrient enriched system would support higher standing crops of fishes in its mangrove zone. We collected 50 species of fish from January 2000 to April 2004 at six sampling sites spanning fresh to brackish salinities in both the Shark and Taylor River drainages. Contrary to expectations, we observed lower standing crops and density of fishes in the more nutrient rich tidal mangrove forest of the Shark River than in the less nutrient rich mangrove habitats bordering the Taylor River. Tidal mangrove habitats in the Shark River were dominated by salt-tolerant fish and displayed lower species richness than mangrove communities in the Taylor River, which included more freshwater taxa and yielded relatively higher richness. These differences were maintained even after controlling for salinity at the time of sampling. Small-scale topographic relief differs between these two systems, possibly created by tidal action in the Shark River. We propose that this difference in topography limits movement of fishes from upstream marshes into the fringing mangrove forest in the Shark River system, but not the Taylor River system. Understanding the influence of habitat structure, including connectivity, on aquatic communities is important to anticipate effects of construction and operational alternatives associated with restoration of the Everglades ecosystem.     

Algal richness and life‐history strategies are influenced by hydrology and phosphorus in two major subtropical wetlands

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Response of Fish Assemblage Structure and Function Following Restoration of Two Small Bahamian Tidal Creeks

Disruption of hydrologic connectivity via road crossings is extremely common in Bahamian tidal creeks, resulting in increased sedimentation and decreased habitat quality and quantity for biota. We restored hydrologic connectivity (i.e., tidal flow) in two small Bahamian mangrove tidal creeks in May 2004 and 2005. We observed the characteristics of fish assemblage structure (species richness) and function (secondary production and transient species utilization of restored areas) before and after restoration, and compared these data with fragmented and unfragmented reference creeks. Restoration significantly increased species richness and secondary production of resident fish species in one of the two restored creeks. Increased utilization of the previously blocked wetlands by transient fishes was observed in both creeks. We suggest success could be attributed to the presence of adjacent nearshore recruitment sources, a more complex local seascape (i.e., high habitat heterogeneity in the creek and local nearshore), and the creation of deep upstream refugia pools. This is one of the first studies to use both structural and functional characteristics to monitor the success of restoration in mangrove ecosystems. Studies combining both structural and functional metrics in restoration monitoring are imperative in linking restoration ecology theory with practical ecological restoration efforts.     

The Southeast Saline Everglades revisited: 50 years of coastal vegetation change

Abstract. We examined the vegetation of the Southeast Saline Everglades (SESE), where water management and sea level rise have been important ecological forces during the last 50 years. Marshes within the SESE were arranged in well‐defined compositional zones parallel to the coast, with mangrove‐dominated shrub communities near the coast giving way to graminoid‐mangrove mixtures, and then Cladium marsh. The compositional gradient was accompanied by an interiorward decrease in total aboveground biomass, and increases in leaf area index and periphyton biomass. Since the mid‐1940s, the boundary of the mixed graminoid‐mangrove and Cladium communities shifted inland by 3.3 km. The interior boundary of a low‐productivity zone appearing white on both black‐and‐white and CIR photos moved inland by 1.5 km on average. A smaller shift in this ‘white zone’ was observed in an area receiving fresh water overflow through gaps in one of the SESE canals, while greater change occurred in areas cut off from upstream water sources by roads or levees. These large‐scale vegetation dynamics are apparently the combined result of sea level rise ‐ ca. 10 cm since 1940 ‐ and water management practices in the SESE.     

Subtroprical wetland fish assemblages and changing salinity regimes: Implications for everglades restoration

During the 1960s, water management practices resulted in the conversion of the wetlands that fringe northeastern Florida Bay (USA) from freshwater/oligohaline herbaceous marshes to dwarf red mangrove forests. Coincident with this conversion were several ecological changes to Florida Bay’s fauna, including reductions in the abundances of top trophic-level consumers: piscivorous fishes, alligators, crocodiles, and wading birds. Because these taxa rely on a common forage base of small demersal fishes, food stress has been implicated as playing a role in their respective declines. In the present study, we monitored the demersal fishes seasonally at six sites over an 8-year time period. During monitoring, extremely high rainfall conditions occurred over a 3.5-year period leading to salinity regimes that can be viewed as “windows” to the area’s natural past and future restored states. In this paper, we: (1) examine the changes in fish communities over the 8-year study period and relate them to measured changes in salinity; (2) make comparisons among marine, brackish and freshwater demersal fish communities in terms of species composition, density, and biomass; and (3) discuss several implications of our findings in light of the intended and unintended water management changes that are planned or underway as part of Everglades restoration. Results suggest the reduction in freshwater flow to Florida Bay over the last several decades has reduced demersal fish populations, and thus prey availability for apex consumers in the coastal wetlands compared to the pre-drainage inferred standard. Furthermore, greater discharge of freshwater toward Florida Bay may result in the re-establishment of pre-1960s fauna, including a more robust demersal-fish community that should prompt increases in populations of several important predatory species.     

Linking stream ecology with morphological variability in a native freshwater fish from semi‐arid Australia

Environmental variation is a potent force affecting phenotypic expression. While freshwater fishes have provided a compelling example of the link between the environment and phenotypic diversity, few studies have been conducted with arid‐zone fishes, particularly those that occur in geographically isolated regions where species typically inhabit intermittent and ephemeral creeks. We investigated morphological variation of a freshwater fish (the western rainbowfish, Melanotaenia australis) inhabiting creeks in the Pilbara region of northwest Australia to determine whether body shape variation correlated with local environmental characteristics, including water velocity, habitat complexity, predator presence, and food availability. We expected that the geographic isolation of creeks within this arid region would result in habitat‐specific morphological specializations. We used landmark‐based geometric morphometrics to quantify the level of morphological variability in fish captured from 14 locations within three distinct subcatchments of a major river system. Western rainbowfish exhibited a range of morphologies, with variation in body depth accounting for a significant proportion (>42%) of the total variance in shape. Sexual dimorphism was also apparent, with males displaying deeper bodies than females. While the measured local habitat characteristics explained little of the observed morphological variation, fish displayed significant morphological differentiation at the level of the subcatchment. Local adaptation may partly explain the geographic patterns of body shape variation, but fine‐scale genetic studies are required to disentangle the effects of genetic differentiation from environmentally determined phenotypic plasticity in body shape. Developing a better understanding of environment–phenotype relationships in species from arid regions will provide important insights into ecological and evolutionary processes in these unique and understudied habitats.     

Wetlands of the Pacific Island region

The wetlands of 21 countries and territories of the Pacific Islands region are reviewed: American Samoa, Cook Islands, Federated States of Micronesia, Fiji, French Polynesia, Guam, Kiribati, Marshall Islands, Nauru, New Caledonia, Niue, Northern Mariana Islands, Palau, Papua New Guinea, Samoa, Solomon Islands, Tokelau, Tonga, Tuvalu, Vanuatu, and Wallis and Futuna. The regions’ wetlands are classified into seven systems: coral reefs, seagrass beds, mangrove swamps, riverine, lacustrine, freshwater swamp forests and marshes. The diversity of species in each of these groups is at near global maxima at the west of the region, with decline towards the east with increasing isolation, and decreasing island size and age. The community structure is unique in each country, and many have endemic species with the habitat isolation that epitomises this island region. There remain, however, some serious gaps in basic inventory, particularly in freshwater biodiversity. Threats to wetlands include introduced freshwater species, loss of wetlands adjacent to urban growth, downstream effects of mining and land clearance, and over-use of mangrove, seagrass and coral reef resources by predominant subsistence economies that remain in this region. Only five countries are signatories to the Ramsar convention on wetlands, and this only recently with seven sites. Wetland managers have identified the need for community education, baseline surveys and monitoring, better legislation and policy for wetland management, and improved capacity of local communities to allow the wise use of their wetlands.     

Does a long‐term shift in Wood Stork diet foreshadow adaptability to human‐induced rapid environmental change?

To preserve biodiversity, identifying at‐risk populations and developing conservation plans to mitigate the effects of human‐induced rapid environmental change (HIREC) are essential. Changes in diet, especially for food‐limited species, can aid in detecting populations being impacted by HIREC, and characterizing the quality, abundance, and temporal and spatial consistency of newly consumed food items may provide insight concerning the likelihood of a species persisting in a changing environment. We used Wood Storks (Mycteria americana) nesting in the Florida Everglades as a model system to study the possible effects of HIREC on a food‐limited population. We compared the diets of Wood Storks in 2013 and 2014 with those reported during the 1970s before major anthropogenic activities affected the Everglades system and prey availability. Wood Storks in our study consumed more large‐bodied sunfish species (Lepomis spp.), fewer native marsh fishes, and more non‐native fish species than during the 1970s. Large sunfish and non‐native fish are relatively rare in the drying pools of Everglades marshes where storks traditionally forage, suggesting that Wood Storks may be using novel foraging habitats such as created wetlands (i.e., canals and stormwater ponds). Although created wetlands have long hydroperiods conducive to maintaining large‐bodied fishes and could provide alternative foraging habitat when prey availability is reduced in natural marshes, additional studies are needed to determine the extent to which these wetlands are used by Wood Storks and, importantly, the quality of prey items potentially available to foraging Wood Storks in created wetlands.     

Benefits of adjacent habitat patches to the distribution of a crayfish population in a hydro-dynamic wetland landscape

Aquatic macrophyte patches are natural features of wetland ecosystems that serve as habitat for aquatic animals. Previous studies suggest animal densities in Everglades, USA, wetlands are generally less numerous in sawgrass ridges than in deeper lily sloughs. We studied the density distribution of a population of Procambarus fallax in ridge and slough habitat types over a 2-year period, spanning two wet–dry cycles and estimated growth and survival rates under flooded conditions to understand comparative value of each to the crayfish population. Procambarus fallax individuals inhabited and recruited in both marsh habitats. During periods of high water, crayfish densities were similar in both habitats; however, densities in both habitats varied seasonally, leading us to postulate some degree of population redistribution in response to fluctuating water depths. Analysis of size distributions over time revealed juveniles in both habitats and two major recruitment periods each year; distinct juvenile cohorts were present in early winter (Nov–Dec) and mid-summer (July–Aug). An in situ experiment of juvenile growth demonstrated that slough habitat type supported faster growth over ridge habitat. To understand habitat-specific mortality risk, a tethering study during flooded conditions indicated that relative predation risk by aquatic predators was greater in sloughs for all sizes and higher for smaller individuals in both habitats. The comparative importance of ridge and slough balances growth potential and survival probability during flooded conditions. This is the first study through time and across both habitat types analyzing the distribution and size structure of P. fallax population in the Everglades.     

Evidence of yellow perch, largemouth bass and pumpkinseed metapopulations in coastal embayments of Lake Ontario

Coastal embayments have been and will continue to be constructed along the northwest shoreline of Lake Ontario to restore and create warmwater fish habitat. However, very little is known about the biological connections among embayments. Using otolith microchemistry on pumpkinseed, largemouth bass and yellow perch collected from three constructed embayments in 2006?C2009, we confirm that these three species of fish each exist in a metapopulation. We find that juvenile pumpkinseed, largemouth bass and yellow perch occupy embayments different from their natal habitat after their first winter, and for at least pumpkinseed, continue to move among embayments after their second winter. We hypothesize that these fishes move among embayments after haphazardly dispersing from their overwintering habitat to the littoral zone each spring. Habitat restoration and remediation efforts in coastal Great Lakes habitats should take a system-based management approach that considers the spatial proximity of embayments, and attempts to create or preserve connected networks.     

Spatial and temporal variability of dissolved organic matter quantity and composition in an oligotrophic subtropical coastal wetland

Dissolved organic matter (DOM) is an essential component of the carbon cycle and a critical driver in controlling variety of biogeochemical and ecological processes in wetlands. The quality of this DOM as it relates to composition and reactivity is directly related to its sources and may vary on temporal and spatial scales. However, large scale, long-term studies of DOM dynamics in wetlands are still scarce in the literature. Here we present a multi-year DOM characterization study for monthly surface water samples collected at 14 sampling stations along two transects within the greater Everglades, a subtropical, oligotrophic, coastal freshwater wetland-mangrove-estuarine ecosystem. In an attempt to assess quantitative and qualitative variations of DOM on both spatial and temporal scales, we determined dissolved organic carbon (DOC) values and DOM optical properties, respectively. DOM quality was assessed using, excitation emission matrix (EEM) fluorescence coupled with parallel factor analysis (PARAFAC). Variations of the PARAFAC components abundance and composition were clearly observed on spatial and seasonal scales. Dry versus wet season DOC concentrations were affected by dry-down and re-wetting processes in the freshwater marshes, while DOM compositional features were controlled by soil and higher plant versus periphyton sources respectively. Peat-soil based freshwater marsh sites could be clearly differentiated from marl-soil based sites based on EEM–PARAFAC data. Freshwater marsh DOM was enriched in higher plant and soil-derived humic-like compounds, compared to estuarine sites which were more controlled by algae- and microbial-derived inputs. DOM from fringe mangrove sites could be differentiated between tidally influenced sites and sites exposed to long inundation periods. As such coastal estuarine sites were significantly controlled by hydrology, while DOM dynamics in Florida Bay were seasonally driven by both primary productivity and hydrology. This study exemplifies the application of long term optical properties monitoring as an effective technique to investigate DOM dynamics in aquatic ecosystems. The work presented here also serves as a pre-restoration condition dataset for DOM in the context of the Comprehensive Everglades Restoration Plan (CERP).     

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.     

Habitat Use by Fishes in Coral Reefs,Seagrass Beds and Mangrove Habitats in the Philippines

Understanding the interconnectivity of organisms among different habitats is a key requirement for generating effective management plans in coastal ecosystems, particularly when determining component habitat structures in marine protected areas. To elucidate the patterns of habitat use by fishes among coral, seagrass, and mangrove habitats, and between natural and transplanted mangroves, visual censuses were conducted semiannually at two sites in the Philippines during September and March 2010–2012. In total, 265 species and 15,930 individuals were recorded. Species richness and abundance of fishes were significantly higher in coral reefs (234 species, 12,306 individuals) than in seagrass (38 species, 1,198 individuals) and mangrove (47 species, 2,426 individuals) habitats. Similarity tests revealed a highly significant difference among the three habitats. Fishes exhibited two different strategies for habitat use, inhabiting either a single (85.6% of recorded species) or several habitats (14.4%). Some fish that utilized multiple habitats, such as Lutjanus monostigma and Parupeneus barberinus, showed possible ontogenetic habitat shifts from mangroves and/or seagrass habitats to coral reefs. Moreover, over 20% of commercial fish species used multiple habitats, highlighting the importance of including different habitat types within marine protected areas to achieve efficient and effective resource management. Neither species richness nor abundance of fishes significantly differed between natural and transplanted mangroves. In addition, 14 fish species were recorded in a 20-year-old transplanted mangrove area, and over 90% of these species used multiple habitats, further demonstrating the key role of transplanted mangroves as a reef fish habitat in this region.     

Predicting diversity of juvenile neotropical fish communities: patch dynamics versus habitat state in floodplain creeks

The species richness of communities should largely depend on habitat variability and/or on habitat state. We evaluated the ability of habitat variability and habitat state to predict the diversity of juvenile neotropical fish communities in creeks of a river floodplain. The young-fish fauna consisted of 73 taxa, and samples were well distributed over a wide range of relevant temporal and spatial habitat variability. We were unable to demonstrate clear patterns of richness in relation to temporal and spatial habitat variability (if habitat state variables were not included), regardless of the temporal variability scale, the grouping of sites (up- and downstream sites differed in temporal variability patterns), taxonomic units or life stages considered. Using stepwise multiple regression, 36% of the variance in species richness was explained for all data, and at best 47% was explained for all taxonomic units at upstream sites using temporal and spatial habitat variability and habitat state (bank length, mean width, mean water level before fishing and/or water turbidity). Using Monte Carlo simulations, we blindly predicted 31% (all data) and at best 37% (all upstream taxa) of the observed variance in species richness from these model types. This limited precision is probably because rare species produced most of the richness patterns in our creeks. The prediction of these rare species is generally difficult for various reasons, and may be a problem in many ecosystem types. Received: 6 July 1998 / Accepted: 16 November 1998     

Trophic behaviour of juvenile reef fishes inhabiting interlinked mangrove–seagrass habitats in offshore mangrove islets

Stable isotope (δ¹³C and δ¹⁵N) and gut content analyses were used to investigate size‐related feeding habits of four reef fishes (the beaugregory Stegastes leucostictus, the french grunt Haemulon flavolineatum, the schoolmaster snapper Lutjanus apodus and the yellowtail snapper Ocyurus chrysurus) inhabiting an offshore (non‐estuarine) mangrove islet off Belize, Central America. Comparisons of isotopic niche space and Schoener diet similarity index suggested a low to moderate degree of niche overlap between fish size groups. The δ¹³C gradient between mangrove and seagrass prey as well as results of Bayesian mixing models revealed that sampled fishes relied mostly on seagrass prey items. Only small and large juveniles of the carnivorous species L. apodus derived a part of their diet from mangroves by targeting mangrove‐associated Grapsidae crabs and fish prey, respectively. Isotopic niche shifts were particularly obvious for carnivorous fishes that ingested larger prey items (Xanthidae crabs and fishes) during their ontogeny. The utilization of mangrove food resources is less than expected and depends on the ecology and life history of the fish species considered. This research highlights that mangrove‐derived carbon contributed relatively little to the diets of four fish taxa from an offshore mangrove islet.     

Hydrologic restoration in a dynamic subtropical mangrove‐to‐marsh ecotone

Extensive hydrologic modifications in coastal regions across the world have occurred to support infrastructure development, altering the function of many coastal wetlands. Wetland restoration success is dependent on the existence of hydrologic regimes that support development of appropriate soils and the growth and persistence of wetland vegetation. In Florida, United States, the Comprehensive Everglades Restoration Program (CERP) seeks to restore, protect, and preserve water resources of the greater Everglades region. Herein we describe vegetation dynamics in a mangrove‐to‐marsh ecotone within the impact area of a CERP hydrologic restoration project currently under development. Vegetation communities are also described for a similar area outside the project area. We found that vegetation shifts within the impact area occurred over a 7‐year period; cover of herbaceous species varied by location, and an 88% increase in the total number of mangrove seedlings was documented. We attribute these shifts to the existing modified hydrologic regime, which is characterized by a low volume of freshwater sheet flow compared with historical conditions (i.e. before modification), as well as increased tidal influence. We also identified a significant trend of decreasing soil surface elevation at the impact area. The CERP restoration project is designed to increase freshwater sheet flow to the impact area. Information from our study characterizing existing vegetation dynamics prior to implementation of the restoration project is required to allow documentation of long‐term project effects on plant community composition and structure within a framework of background variation, thereby allowing assessment of the project's success in restoring critical ecosystem functions.     

Integrating multidirectional connectivity requirements in systematic conservation planning for freshwater systems

Aim Recent efforts to apply the principles of systematic conservation planning to freshwater ecosystems have focused on the special connected nature of these systems as a way to ensure adequacy (long‐term maintenance of biodiversity). Connectivity is important in maintaining biodiversity and key ecological processes in freshwater environments and is of special relevance for conservation planning in these systems. However, freshwater conservation planning has focused on longitudinal connectivity requirements within riverine ecosystems, while other habitats, such as floodplain wetlands or lakes and connections among them, have been overlooked. Here, we address this gap by incorporating a new component of connectivity in addition to the traditional longitudinal measure. Location Northern Australia. Methods We integrate lateral connections between freshwater areas (e.g. lakes and wetlands) that are not directly connected by the river network and the longitudinal upstream–downstream connections. We demonstrate how this can be used to incorporate ecological requirements of some water‐dependent taxa that can move across drainage divides, such as waterbirds. Results When applied together, the different connectivity rules allow the identification of priority areas that contain whole lakes or wetlands, their closest neighbours whenever possible, and the upstream/downstream reaches of rivers that flow into or from them. This would facilitate longitudinal and lateral movements of biota while minimizing the influence of disturbances potentially received from upstream or downstream reaches. Main conclusions This new approach to defining and applying different connectivity rules can help improve the adequacy of freshwater‐protected areas by enhancing movements of biodiversity within priority areas. The integration of multiple connectivity needs can also serve as a bridge to integrate freshwater and terrestrial conservation planning.     

Annual and non-annual fish assemblages respond differently to environmental and spatiotemporal variations of temporary wetlands from southern Brazil

• The fish community of temporary wetlands is highly dynamic due to the marked seasonal changes in hydrological and limnological conditions. In this study, we evaluated spatiotemporal variation in the fish community and the influence of environmental factors on this variation in temporary wetlands in southern Brazil.
• Two different life cycles—annual and non-annual fishes—were recorded among species. Variation in the fish community appeared to be associated with differences in the life history strategy of species. While species of annual fish were present throughout the hydroperiod, gradually decreasing in density over time, non-annual fish species had an opposite pattern, with higher densities in the final phase of the hydrological cycle.
• Variations in the composition and abundance of fish community in temporary wetlands were related to habitat and landscape attributes (area, number of microhabitats and isolation between temporary wetlands) and climate indicators of water availability (air temperature, average rainfall, and water level of the São Gonçalo channel), but not to water conditions (dissolved oxygen, pH, conductivity and water temperature).
• We concluded that the ichthyofauna of temporary wetlands are composed both of rare species, highly specialised for life within these ephemeral environments, and widespread generalist species that reach the wetlands opportunistically for short periods. Habitat and landscape attributes and changes in climate and hydrological variables over the hydroperiod are the factors that had the greatest influence on spatiotemporal variation of the fish community.
• Maintaining physical characteristics and the hydrological cycle of temporary wetlands is critical for the conservation of fish species inhabiting these vulnerable environments in southern Brazil. Habitat and landscape attributes and climate and hydrology fluctuations affect the annual and non-annual fishes inhabiting temporary wetlands differently.