Diel Patterns of Patch Reef Community Dynamics in a Caribbean Back-Reef System

Back-reef seascapes represent critical habitat for juvenile and adult fishes. Patch reef, seagrass, and mangrove habitats form a heterogeneous mosaic, often linked by species that use reefs as structure during the day and make foraging migrations into soft-bottom habitat at night. Artificial reefs are used to model natural patch reefs, however may not function equivalently as fish habitat. To study the relative value of natural and artificial patch reefs as fish habitat, these communities in the Sea of Abaco, Bahamas were compared using roving diver surveys and time-lapse photography. Diel turnover in fish abundance, recorded with time-lapse photography and illuminated by infrared light, was quantified across midday, dusk, and night periods to explore possible effects of reef type (artificial vs. natural) on these patterns. Diurnal communities on natural reefs exhibited greater fish abundance, species richness, and functional diversity compared to artificial reefs. Furthermore, both types of reef communities exhibited a significant shift across the diel period, characterized by a decline in total fish density at night, especially for grunts (Haemulidae). Cross-habitat foraging migrations by diurnal or nocturnal species, such as haemulids, are likely central drivers of this twilight turnover and can represent important energy and nutrient subsidies. Time-lapse surveys provided more consistent measures of reef fish assemblages for the smaller artificial reef habitats, yet underestimated abundance of certain taxa and species richness on larger patch habitats when compared to the roving diver surveys. Time-lapse photography complemented with infrared light represent a valuable non-invasive approach to studying behavior of focal species and their fine-scale temporal dynamics in shallow-reef communities.

Spatial scale and abundance patterns of large fish communities in freshwater marshes of the Florida Everglades

Anthropogenic habitat alterations and water-management practices have imposed an artificial spatial scale onto the once contiguous freshwater marshes of the Florida Everglades. To gain insight into how these changes may affect biotic communities, we examined whether variation in the abundance and community structure of large fishes (SL . 8 cm) in Everglades marshes varied more at regional or intraregional scales, and whether this variation was related to hydroperiod, water depth, floating mat volume, and vegetation density. From October 1997 to October 2002, we used an airboat electrofisher to sample large fishes at sites within three regions of the Everglades. Each of these regions is subject to unique watermanagement schedules. Dry-down events (water depth , 10 cm) occurred at several sites during spring in 1999, 2000, 2001, and 2002. The 2001 dry-down event was the most severe and widespread. Abundance of several fishes decreased significantly through time, and the number of days post-dry-down covaried significantly with abundance for several species. Processes operating at the regional scale appear to play important roles in regulating large fishes. The most pronounced patterns in abundance and community structure occurred at the regional scale, and the effect size for region was greater than the effect size for sites nested within region for abundance of all species combined, all predators combined, and each of the seven most abundant species. Non-metric multi-dimensional scaling revealed distinct groupings of sites corresponding to the three regions. We also found significant variation in community structure through time that correlated with the number of days post-dry-down. Our results suggest that hydroperiod and water management at the regional scale influence large fish communities of Everglades marshes.

The influence of body size on the ecology of coastal fish predators in the Bahamas

Body size is a fundamental structural characteristic of organisms, determining critical life history and physiological traits, and influencing population dynamics, community structure, and ecosystem function. For my dissertation, I focused on effects of body size on habitat use and diet of important coastal fish predators, as well as their influence on faunal communities in Bahamian wetlands. First, using acoustic telemetry and stable isotope analysis, I identified high variability in movement patterns and habitat use among individuals within a gray snapper (Lutjanus griseus) and schoolmaster snapper (L. apodus) population. This intrapopulation variation was not explained by body size, but by individual behavior in habitat use. Isotope values differed between individuals that moved further distances and individuals that stayed close to their home sites, suggesting movement differences were related to specific patterns of foraging behavior. Subsequently, while investigating diet of schoolmaster snapper over a two-year period using stomach content and stable isotope analyses, I also found intrapopulation diet variation, mostly explained by differences in size class, individual behavior and temporal variability. I then developed a hypothesis-testing framework examining intrapopulation niche variation between size classes using stable isotopes. This framework can serve as baseline to categorize taxonomic or functional groupings into specific niche shift scenarios, as well as to help elucidate underlying mechanisms causing niche shifts in certain size classes. Finally, I examined the effect of different-sized fish predators on epifaunal community structure in shallow seagrass beds using exclusion experiments at two spatial scales. Overall, I found that predator effects were rather weak, with predator size and spatial scale having no impact on the community. Yet, I also found some evidence of strong interactions on particular common snapper prey. As Bahamian wetlands are increasingly threatened by human activities (e.g., overexploitation, habitat degradation), an enhanced knowledge of the ecology of organisms inhabiting these systems is crucial for developing appropriate conservation and management strategies. My dissertation research contributed to this effort by providing critical information about the resource use of important Bahamian fish predators, as well as their effect on faunal seagrass communities.

Canals as Vectors for Fish Movement: Potential Southward Range Expansion of Lepisosteus osseus L. (Longnose Gar) in South Florida

Lepisosteus osseus (Longnose Gar) is a large-bodied predator, whose Florida distribution remains unclear at the southern edge of its range. We reviewed available literature and museum voucher specimens to provide a more accurate range description, and we discuss recent collections in south Florida. Longnose Gar has not been previously reported in natural habitats south of Lake Okeechobee. Instead, records south of the lake are from canals, and most are recent (since 2000), including our own southernmost 2011 record. No records from Everglades natural habitats have been collected. Previous studies have shown native range expansions in anthropogenically disturbed landscapes. We suggest that the Longnose Gar is expanding its range southward in Florida using canals as dispersal vectors and/or suitable habitat.

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.

Using data from an encounter sampler to model fish dispersal

A method to estimate speed of free-ranging fishes using a passive sampling device is described and illustrated with data from the Everglades, U.S.A. Catch per unit effort (CPUE) from minnow traps embedded in drift fences was treated as an encounter rate and used to estimate speed, when combined with an independent estimate of density obtained by use of throw traps that enclose 1 m2 of marsh habitat. Underwater video was used to evaluate capture efficiency and species-specific bias of minnow traps and two sampling studies were used to estimate trap saturation and diel-movement patterns; these results were used to optimize sampling and derive correction factors to adjust species-specific encounter rates for bias and capture efficiency. Sailfin mollies Poecilia latipinna displayed a high frequency of escape from traps, whereas eastern mosquitofish Gambusia holbrooki were most likely to avoid a trap once they encountered it; dollar sunfish Lepomis marginatus were least likely to avoid the trap once they encountered it or to escape once they were captured. Length of sampling and time of day affected CPUE; fishes generally had a very low retention rate over a 24 h sample time and only the Everglades pygmy sunfish Elassoma evergladei were commonly captured at night. Dispersal speed of fishes in the Florida Everglades, U.S.A., was shown to vary seasonally and among species, ranging from 0· 05 to 0· 15 m s−1 for small poeciliids and fundulids to 0· 1 to 1· 8 m s−1 for L. marginatus. Speed was generally highest late in the wet season and lowest in the dry season, possibly tied to dispersal behaviours linked to finding and remaining in dry-season refuges. These speed estimates can be used to estimate the diffusive movement rate, which is commonly employed in spatial ecological models.

Hydroperiod and seasonal effects on fish decomposition in an oligotrophic Everglades marsh

In the Everglades, the majority of fish detrital inputs occur during the dry scason, when waterlevel drawdown reduces aquatic habitat. While these mortality events are highly seasonal, the remineralization and recycling of fish detrital nutrients may represent an important stimulus to the ecosystem in the following wet season. The goal of this study was to quantify the rate of detrital fish decomposition during three periods of the year to determine seasonal variations in decomposition patterns in this ecosystem. A multiple regression analysis showed that hydroperiod and water depth both played a role in determining fish decomposition rates within this ecosystem. Decomposition rates ranged from a low of 13% day−1 in December 2000 to a high of 50% day−1 in April 2001, the height of the dry season. Phosphorus analysis showed that Gambusia holbrooki, the dominant small fish species in the Everglades, contains 7.169±1.46 mg P g−1 wet fish weight. Based on the observed decomposition rates and the average biomass added, the estimafed daily flux of phosphorus from the experimental detrital loading ranged from a low of 27.04 mg P day−1 to a high of 108.14 mg P day−1 during the decomposition period. We estimated that these inputs could represent an input of 43 μg P m−2 day−1 to the total temporal Everglades phosphorus budget. Although much of this phosphorus is likely incorporated into the macroinvertebrate pool, detrital inputs peak during the dry season when nutrients are most likely to be incorporated into the soil and occur when decomposition of vegetative material is moisture-limited. These inputs may therefore play an important role in stimulating vegetative production during the early wet season.

Does landscape context affect habitat value? The importance of seascape ecology in back-reef systems

Seascape ecology provides a useful framework from which to understand the processes governing spatial variability in ecological patterns. Seascape context, or the composition and pattern of habitat surrounding a focal patch, has the potential to impact resource availability, predator-prey interactions, and connectivity with other habitats. For my dissertation research, I combined a variety of approaches to examine how habitat quality for fishes is influenced by a diverse range of seascape factors in sub-tropical, back-reef ecosystems. In the first part of my dissertation, I examined how seascape context can affect reef fish communities on an experimental array of artificial reefs created in various seascape contexts in Abaco, Bahamas. I found that the amount of seagrass at large spatial scales was an important predictor of community assembly on these reefs. Additionally, seascape context had differing effects on various aspects of habitat quality for the most common reef species, White grunt Haemulon plumierii. The amount of seagrass at large spatial scales had positive effects on fish abundance and secondary production, but not on metrics of condition and growth. The second part of my dissertation focused on how foraging conditions for fish varied across a linear seascape gradient in the Loxahatchee River estuary in Florida, USA. Gray snapper, Lutjanus griseus, traded food quality for quantity along this estuarine gradient, maintaining similar growth rates and condition among sites. Additional work focused on identifying major energy flow pathways to two consumers in oyster-reef food webs in the Loxahatchee. Algal and microphytobenthos resource pools supported most of the production to these consumers, and body size for one of the consumers mediated food web linkages with surrounding mangrove habitats. All of these studies examined a different facet of the importance of seascape context in governing ecological processes occurring in focal habitats and underscore the role of connectivity among habitats in back-reef systems. The results suggest that management approaches consider the surrounding seascape when prioritizing areas for conservation or attempting to understand the impacts of seascape change on focal habitat patches. For this reason, spatially-based management approaches are recommended to most effectively manage back-reef systems.

Characterization of Atlantic Cod Spawning Habitat and Behavior in Icelandic Coastal Waters

The physical habitat used during spawning may potentially be an important factor affecting reproductive output of broadcast spawning marine fishes, particularly for species with complex, substrate-oriented mating systems and behaviors, such as Atlantic cod Gadus morhua. We characterized the habitat use and behavior of spawning Atlantic cod at two locations off the coast of southwestern Iceland during a 2-d research cruise (15–16 April 2009). We simultaneously operated two different active hydroacoustic gear types, a split beam echosounder and a dual frequency imaging sonar (DIDSON), as well as a remotely operated underwater vehicle (ROV). A total of five fish species were identified through ROV surveys: including cusk Brosme brosme, Atlantic cod, haddock Melanogrammus aeglefinus, lemon sole Microstomus kitt, and Atlantic redfish Sebastes spp. Of the three habitats identified in the acoustic surveys, the transitional habitat between boulder/lava field and sand habitats was characterized by greater fish density and acoustic target strength compared to that of sand or boulder/lava field habitats independently. Atlantic cod were observed behaving in a manner consistent with published descriptions of spawning. Individuals were observed ascending 1–5 m into the water column from the bottom at an average vertical swimming speed of 0.20–0.25 m s−1 and maintained an average spacing of 1.0–1.4 m between individuals. Our results suggest that cod do not choose spawning locations indiscriminately despite the fact that it is a broadcast spawning fish with planktonic eggs that are released well above the seafloor.

Turning Passive Detection Systems into Field Experiments: An Application Using Wetland Fishes and Enclosures to Track Fine-Scale Movement and Habitat Choice

Understanding habitat selection and movement remains a key question in behavioral ecology. Yet, obtaining a sufficiently high spatiotemporal resolution of the movement paths of organisms remains a major challenge, despite recent technological advances. Observing fine-scale movement and habitat choice decisions in the field can prove to be difficult and expensive, particularly in expansive habitats such as wetlands. We describe the application of passive integrated transponder (PIT) systems to field enclosures for tracking detailed fish behaviors in an experimental setting. PIT systems have been applied to habitats with clear passageways, at fixed locations or in controlled laboratory and mesocosm settings, but their use in unconfined habitats and field-based experimental setups remains limited. In an Everglades enclosure, we continuously tracked the movement and habitat use of PIT-tagged centrarchids across three habitats of varying depth and complexity using multiple flatbed antennas for 14 days. Fish used all three habitats, with marked species-specific diel movement patterns across habitats, and short-lived movements that would be likely missed by other tracking techniques. Findings suggest that the application of PIT systems to field enclosures can be an insightful approach for gaining continuous, undisturbed and detailed movement data in unconfined habitats, and for experimentally manipulating both internal and external drivers of these behaviors.

Seasonal Fish Dispersal in Ephemeral Wetlands of the Florida Everglades

We hypothesized that fishes in short-hydroperiod wetlands display pulses in activity tied to seasonal flooding and drying, with relatively low activity during intervening periods. To evaluate this hypothesis, sampling devices that funnel fish into traps (drift fences) were used to investigate fish movement across the Everglades, U.S.A. Samples were collected at six sites in the Rocky Glades, a seasonally flooded karstic habitat located on the southeastern edge of the Everglades. Four species that display distinct recovery patterns following drought in long-hydroperiod wetlands were studied: eastern mosquitofish (Gambusia holbrooki) and flagfish (Jordanella floridae) (rapid recovery); and bluefin killifish (Lucania goodei) and least killifish (Heterandria formosa) (slow recovery). Consistent with our hypothesized conceptual model, fishes increased movement soon after flooding (immigration period) and just before drying (emigration period), but decreased activity in the intervening foraging period. We also found that eastern mosquitofish and flagfish arrived earlier and showed stronger responses to hydrological variation than either least killifish or bluefin killifish. We concluded that these fishes actively colonize and escape ephemeral wetlands in response to flooding and drying, and display species-specific differences related to flooding and drying that reflect differences in dispersal ability. These results have important implications for Everglades fish metacommunity dynamics.

Effect of Hydrologic Restoration on the Habitat of The Cape Sable Seaside Sparrow, Annual Report of 2004-2005

The major activities in Year 3 on ‘Effect of hydrologic restoration on the habitat of the Cape Sable seaside sparrow (CSSS)’ included presentations, field work, data analysis, and report preparation. During this period, we made 4 presentations, two at the CSSS – fire planning workshops at Everglades National Park (ENP), one at the Society of Wetland Scientists’ meeting in Charleston, SC, and a fourth at the Marl Prairie/CSSS performance measure workshop at ENP. We started field work in the third week of January and continued till June 3, 2005. Early in the field season, we completed vegetation surveys along two transects, B and C (~15.1 km). During April and May, vegetation sampling was completed at 199 census sites, bringing to 608 the total number of CSSS census sites with quantitative vegetation data. We updated data sets from all three years, 2003-05, and analyzed them using cluster analysis and ordination as in previous two years. However, instead of weighted averaging, we used weighted-averaging partial least square regression (WA-PLS) model, as this method is considered an improvement over WA for inferring values of environmental variables from biological species composition. We also validated the predictive power of the WA-PLS regression model by applying it to a sub-set of 100 census sites for which hydroperiods were “known” from two sources, i.e., from elevations calculated from concurrent water depth measurements onsite and at nearby water level recorders, and from USGS digital elevation data. Additionally, we collected biomass samples at 88 census sites, and determined live and dead aboveground plant biomass. Using vegetation structure and biomass data from those sites, we developed a regression model that we used to predict aboveground biomass at all transects and census sites. Finally, biomass data was analyzed in relation to hydroperiod and fire frequency.

Woody Plant Invasion into the Freshwater Marl Prairie Habitat of the Cape Sable Seaside Sparrow: Final Report

In the fall of 2005, U.S. Fish and Wildlife Services (USFWS) contracted with Florida International University (FIU) to study the physical and biological drivers underlying the distribution of woody plant species in the marl prairie habitat of the Cape Sable Seaside Sparrow (CSSS). This report presents what we have learned about woody plant encroachment based on studies carried out during the period 2006-2008. The freshwater marl prairie habitat currently occupied by the Cape Sable seaside sparrow (CSSS; Ammodramus maritimus mirabilis) is a dynamic mosaic comprised of species-rich grassland communities and tree islands of various sizes, densities and compositions. Landscape heterogeneity and the scale of vegetative components across the marl prairie is primarily determined by hydrologic conditions, biological factors (e.g. dispersal and growth morphology), and disturbances such as fire. The woody component of the marl prairie landscape is subject to expansion through multiple positive feedback mechanisms, which may be initiated by recent land use change (e.g. drainage). Because sparrows are known to avoid areas where the woody component is too extensive, a better understanding of invasion dynamics is needed to ensure proper management.

Does Landscape Context Affect Habitat Value? The Importance of Seascape Ecology in Back-reef Systems

Seascape ecology provides a useful framework from which to understand the processes governing spatial variability in ecological patterns. Seascape context, or the composition and pattern of habitat surrounding a focal patch, has the potential to impact resource availability, predator-prey interactions, and connectivity with other habitats. For my dissertation research, I combined a variety of approaches to examine how habitat quality for fishes is influenced by a diverse range of seascape factors in sub-tropical, back-reef ecosystems. In the first part of my dissertation, I examined how seascape context can affect reef fish communities on an experimental array of artificial reefs created in various seascape contexts in Abaco, Bahamas. I found that the amount of seagrass at large spatial scales was an important predictor of community assembly on these reefs. Additionally, seascape context had differing effects on various aspects of habitat quality for the most common reef species, White grunt Haemulon plumierii. The amount of seagrass at large spatial scales had positive effects on fish abundance and secondary production, but not on metrics of condition and growth. The second part of my dissertation focused on how foraging conditions for fish varied across a linear seascape gradient in the Loxahatchee River estuary in Florida, USA. Gray snapper, Lutjanus griseus, traded food quality for quantity along this estuarine gradient, maintaining similar growth rates and condition among sites. Additional work focused on identifying major energy flow pathways to two consumers in oyster-reef food webs in the Loxahatchee. Algal and microphytobenthos resource pools supported most of the production to these consumers, and body size for one of the consumers mediated food web linkages with surrounding mangrove habitats. All of these studies examined a different facet of the importance of seascape context in governing ecological processes occurring in focal habitats and underscore the role of connectivity among habitats in back-reef systems. The results suggest that management approaches consider the surrounding seascape when prioritizing areas for conservation or attempting to understand the impacts of seascape change on focal habitat patches. For this reason, spatially-based management approaches are recommended to most effectively manage back-reef systems.