Comparative reproductive biology of two Florida pawpaws Asimina reticulata Chapman and Asimina tetramera Small

I investigated the phenology and breeding systems of two Florida endemic pawpaws, Asimina reticulata, widespread in peninsular Florida, and A. tetramera, a federally endangered species limited to two counties on the Atlantic Coastal Ridge. The purpose of this study was to determine if differences contribute to the rarity of Asimina tetramera compared with A. reticulata. The study was conducted in sand pine scrub sites with the largest populations of A. tetramera in the two counties. Flowering seasons differ for the two species. Both species are hermaphroditic and strongly protogynous. Pollination experiments show that neither species is autogamous and the primary breeding mechanism is outcrossing, although low levels of geitonogamous pollination occur in mature scrub habitats. High levels of inbreeding depression were noted in both species at both sites but inbreeding depression was relaxed the first year post-fire. Fruit set in mature habitats may be pollinator limited. ^ I studied insects associated with the flowers in sand pine scrub habitat in southeastern Florida from 1994–1996. The most commonly represented orders were Coleoptera (25 spp.), Lepidoptera. (3 spp.) and Hymenoptera. (3 spp.). All Coleoptera. were flower visitors; one species, Euphoria sepulchralis (Fabricius)(Scarabeaidae), visited flowers of the two Asimina species at both sites. Eurytides marcellus (Cramer) (Lepidoptera: Papilionidae) eggs and larvae were observed on both species of Asimina during each year of the study. ^ Resource management techniques were applied to a mature sand pine scrub community in Jonathan Dickinson State Park in southeastern Florida for the management of Asimina tetramera. Manipulations conducted in 1996 included combinations of fire and mechanical treatments. I measured effects of these treatments on flowering and fruit set on A. tetramera and found cutting and burning was most effective in increasing flowering, followed by burning. Mechanical cutting and mulching had no significant effect. ^

Conservation implications of the reproductive biology of the endangered vine Ipomoea microdactyla Griseb. (Convolvulaceae)

A plant's reproductive biology exerts a significant influence on both population persistence within changing environments and successful establishment of new populations. However, the interaction between extrinsic (i.e. ecological) and intrinsic (i.e. genetic) factors also is an important driver of demographic performance for plant populations. It is light of this that I performed a multidisciplinary investigation of the breeding system, seed and seedling establishment dynamics, and population genetic structure of the endangered Caribbean vine Ipomoea microdactyla Griseb. (Convolvulaceae). The results from the breeding system study show individuals from Florida, USA and Andros Island, Bahamas to be self-incompatible. Plants from the two regions are cross-compatible but there is evidence for outbreeding depression in their progeny. Significant regional differences were found in floral traits and progeny traits that suggests incipient speciation for the Florida populations. The results from the seed and seedling establishment dynamics experiment demonstrate that the restoration of small populations in Florida via seed and seedling augmentation is a successful strategy. The demographic performance of the outplanted individuals was driven significantly by ecological factors (e.g. herbivory) rather than by genetic factors which emphasizes that the ecological context is very important for successful restoration attempts. The results from the population genetic study using an analysis of molecular variation (AMOVA) reveal significant differences in genetic variation among individuals from Florida, Andros, and Cuba. A Bayesian analysis of population genetic structuring coincided with the previous AMOVA results among the three regions. The Mantel test indicated significant 'isolation by distance' for these regional populations implying restricted gene flow over relatively short distances. Overall, the Florida populations had the lowest measures of genetic diversity which is most likely due to the effects of both colonization founder events and habitat fragmentation. The results of my study highlight the value of performing multidisciplinary studies in relation to species conservation as knowledge of both extrinsic and intrinsic factors can best guide decisions for species preservation.

Influence of habitat on the reproductive ecology of the Amazonian palm, Mauritia flexuosa, in Roraima, Brazil

Although Mauritia flexuosa (Arecaceae) plays a pivotal role in the ecology and economy of the Amazon, and occurs in a variety of habitats, little is known about the influence of habitat on the reproductive biology of this palm. My dissertation focuses on the reproductive biology of M. flexuosa in three habitats in Roraima, Brazil: undisturbed forest, undisturbed forest-savanna ecotone, and savanna disturbed by plantations of the exotic tree, Acacia mangium. First, I calculated sex ratios and linked precipitation patterns with phenology. Sex ratios were female-biased. Precipitation was negatively associated with flowering, and positively associated with fruiting. Habitat appears to have no significant influence on phenology of M. flexuosa, although short-term climate variation may affect phenology of this species. Second, I examined floral biology, observed floral visitors, and performed exclusion experiments to determine the pollination system of M. flexuosa. Fruit set did not differ significantly between the visitor exclusion treatment and the control, but was significantly lowest in the wind + visitor exclusion treatment, suggesting that this dioecious palm is anemophilous, independent of habitat. Third, I identified the abiotic and biotic factors explaining variation in fruit mass, seed mass, seed number per fruit, and total fruit yield among habitats. Soil moisture and flooding during the wet season were the best predictors of fruit and seed output. The number of leaves, diameter at breast height, and height were all accurate predictors of reproductive output, but crown volume did not accurately predict fruit yields. Results re-evaluate traditional assumptions about wind-pollination in the tropics, and highlight abiotic and biotic factors responsible for variation in reproductive output of M. flexuosa, with implications for effective management of this palm. Finally, I interviewed harvesters and vendors to document the traditional knowledge and market dynamics of the fruit of M. flexuosa, buriti. Traditional knowledge corroborated results from scientific studies. Vendors argued that the price of buriti must increase, and must fluctuate with varying supply. With appropriate economic incentives to vendors/harvesters, Roraima may expand its market infrastructure for buriti, effectively stimulating the regional economy and practicing sustainable harvesting.

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.

Microhabitat Selection by Marine Mesoconsumers in a Thermally Heterogeneous Habitat: Behavioral Thermoregulation or Avoiding Predation Risk?

Habitat selection decisions by consumers has the potential to shape ecosystems. Understanding the factors that influence habitat selection is therefore critical to understanding ecosystem function. This is especially true of mesoconsumers because they provide the link between upper and lower tropic levels. We examined the factors influencing microhabitat selection of marine mesoconsumers – juvenile giant shovelnose rays (Glaucostegus typus), reticulate whiprays (Himantura uarnak), and pink whiprays (H. fai) – in a coastal ecosystem with intact predator and prey populations and marked spatial and temporal thermal heterogeneity. Using a combination of belt transects and data on water temperature, tidal height, prey abundance, predator abundance and ray behavior, we found that giant shovelnose rays and reticulate whiprays were most often found resting in nearshore microhabitats, especially at low tidal heights during the warm season. Microhabitat selection did not match predictions derived from distributions of prey. Although at a course scale, ray distributions appeared to match predictions of behavioral thermoregulation theory, fine-scale examination revealed a mismatch. The selection of the shallow nearshore microhabitat at low tidal heights during periods of high predator abundance (warm season) suggests that this microhabitat may serve as a refuge, although it may come with metabolic costs due to higher temperatures. The results of this study highlight the importance of predators in the habitat selection decisions of mesoconsumers and that within thermal gradients, factors, such as predation risk, must be considered in addition to behavioral thermoregulation to explain habitat selection decisions. Furthermore, increasing water temperatures predicted by climate change may result in complex trade-offs that might have important implications for ecosystem dynamics.

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.

Pollination, Herbivory, and Habitat Fragmentation: Their Effects on the Reproductive Fitness of Angadenia berteroi, a Native Perennial Plant of the South Florida Pine Rocklands

Angadenia berteroi is a tropical perennial subshrub of the pine rocklands with large yellow flowers that set very few fruits. My dissertation seeks to elucidate the factors that affect the reproductive fitness of Angadenia berteroi a native species of the south Florida pine rocklands. I provide novel information on the pollination biology of this native species. I also assess the effects of herbivory on growth and the reproductive success of A. berteroi. Finally, I elucidate how habitat fragmentation and quality are correlated with reproductive fitness of this native perennial plant. Using a novel experimental approach, I determined the most effective pollinator group. I used nylon fishing line of widths corresponding to proboscis diameter of the major groups of visitors to examine pollen removal and deposition. In the field, I estimated visitation frequency and efficacy of each pollinator type. Using potted plants, I exposed flowers to single visit from different types of pollinators to measure fruit set. I performed artificial defoliation with scissors on plants growing in the greenhouse to assess the effects of defoliation before flowering as well as during flowering. Additionally, I used structural equation modelling (SEM) to elucidate how A. berteroi reproductive fitness was affected by habitat fragmentation and quality. My experiments provide evidence that Angadenia berteroi is specialized for bee pollination; though butterflies, skippers and others also visit its flowers, A. berteroi is exclusively pollinated by two native bees of the South Florida pine rocklands . This research also demonstrated that herbivory by the oleander moth may have direct and indirect effects on Angadenia berteroi growth and reproductive success. The SEM results suggested that habitat quality (litter depth and subcanopy cover) may favor reproduction in native species of the South Florida pine rocklands that are properly maintained by periodic fires and exotic control. Insights from this threatened and charismatic species may provide impetus to properly manage remaining pine rocklands in South Florida for this and other endemic understory species.

The rodent fauna of Long Pine Key, Everglades National Park: a comparison of habitat types

Rodents are often involved at several stages of trophic dynamics. Consequently they often play crucial roles in the structure and function of many complex ecological systems. This study sought to address the lack of baseline data concerning rodents in tropical areas, and south Florida in particular. Live trapping took place in the four major habitat types of the Long Pine Key area of Everglades National Park over the course of one year. I compared population structures and abundance of murid rodents in the four habitat types, and tested multiple weather variables for their effectiveness as predictors of rodent abundance. I found the Long Pine Key area to be depauperate in terms of species diversity. Each of the four species of rodent encountered favored a particular habitat type. The density of the understory vegetation and the avoidance of avian predators in particular appear to be the most important factors in the distribution and abundance of rodents in the Long Pine Key area of Everglades National Park.

Habitat Use of the Key Largo Woodrat (Neotoma floridana smalli)

Key Largo woodrats are an endangered subspecies with an extremely limited habitat. This study sought to understand woodrat habitat preferences in order to guide management. Woodrats build stick nests from natural and artificial materials, so nest distribution and nest occupancy were used as indicators of preference. Distribution was determined by nest surveys, and remote cameras were used to assess occupancy. Forest structure, human disturbance, nest, and animal presence metrics were also collected. More nests were found along abandoned roads than along forest transects and more artificial nests were occupied than natural nests. These findings indicate that woodrats prefer areas with human disturbance, rather than forest age and structure as previously believed. This may have consequences on woodrat management, as it suggests that man-made materials are currently needed by woodrats even in a protected natural area.

Influence of Habitat on the Reproductive Ecology of the Amazonian Palm, Mauritia flexuosa, in Roraima, Brazil

Although Mauritia flexuosa (Arecaceae) plays a pivotal role in the ecology and economy of the Amazon, and occurs in a variety of habitats, little is known about the influence of habitat on the reproductive biology of this palm. My dissertation focuses on the reproductive biology of M. flexuosa in three habitats in Roraima, Brazil: undisturbed forest, undisturbed forest-savanna ecotone, and savanna disturbed by plantations of the exotic tree, Acacia mangium. First, I calculated sex ratios and linked precipitation patterns with phenology. Sex ratios were female-biased. Precipitation was negatively associated with flowering, and positively associated with fruiting. Habitat appears to have no significant influence on phenology of M. flexuosa, although short-term climate variation may affect phenology of this species. Second, I examined floral biology, observed floral visitors, and performed exclusion experiments to determine the pollination system of M. flexuosa. Fruit set did not differ significantly between the visitor exclusion treatment and the control, but was significantly lowest in the wind + visitor exclusion treatment, suggesting that this dioecious palm is anemophilous, independent of habitat. Third, I identified the abiotic and biotic factors explaining variation in fruit mass, seed mass, seed number per fruit, and total fruit yield among habitats. Soil moisture and flooding during the wet season were the best predictors of fruit and seed output. The number of leaves, diameter at breast height, and height were all accurate predictors of reproductive output, but crown volume did not accurately predict fruit yields. Results re-evaluate traditional assumptions about wind-pollination in the tropics, and highlight abiotic and biotic factors responsible for variation in reproductive output of M. flexuosa, with implications for effective management of this palm. Finally, I interviewed harvesters and vendors to document the traditional knowledge and market dynamics of the fruit of M. flexuosa, buriti. Traditional knowledge corroborated results from scientific studies. Vendors argued that the price of buriti must increase, and must fluctuate with varying supply. With appropriate economic incentives to vendors/harvesters, Roraima may expand its market infrastructure for buriti, effectively stimulating the regional economy and practicing sustainable harvesting.

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.

Pollination, herbivory, and habitat fragmentation: Their effects on the reproductive fitness of Angadenia berteroi, a native perennial plant of the south Florida pine rocklands

Angadenia berteroi is a tropical perennial subshrub of the pine rocklands with large yellow flowers that set very few fruits. My dissertation seeks to elucidate the factors that affect the reproductive fitness of Angadenia berteroi a native species of the south Florida pine rocklands. I provide novel information on the pollination biology of this native species. I also assess the effects of herbivory on growth and the reproductive success of A. berteroi. Finally, I elucidate how habitat fragmentation and quality are correlated with reproductive fitness of this native perennial plant.^ Using a novel experimental approach, I determined the most effective pollinator group. I used nylon fishing line of widths corresponding to proboscis diameter of the major groups of visitors to examine pollen removal and deposition. In the field, I estimated visitation frequency and efficacy of each pollinator type. Using potted plants, I exposed flowers to single visit from different types of pollinators to measure fruit set. I performed artificial defoliation with scissors on plants growing in the greenhouse to assess the effects of defoliation before flowering as well as during flowering. Additionally, I used structural equation modelling (SEM) to elucidate how A. berteroi reproductive fitness was affected by habitat fragmentation and quality. ^ My experiments provide evidence that Angadenia berteroi is specialized for bee pollination; though butterflies, skippers and others also visit its flowers, A. berteroi is exclusively pollinated by two native bees of the South Florida pine rocklands . This research also demonstrated that herbivory by the oleander moth may have direct and indirect effects on Angadenia berteroi growth and reproductive success. The SEM results suggested that habitat quality (litter depth and subcanopy cover) may favor reproduction in native species of the South Florida pine rocklands that are properly maintained by periodic fires and exotic control. Insights from this threatened and charismatic species may provide impetus to properly manage remaining pine rocklands in South Florida for this and other endemic understory species.^