Breaking down complex saproxylic communities: understanding sub-networks structure and implications to network robustness

Saproxylic insect communities inhabiting tree hollow microhabitats correspond with large food webs which simultaneously are constituted by multiple types of plant-animal and animal-animal interactions, according to the use of trophic resources (wood- and insect-dependent sub-networks), or to trophic habits or interaction types (xylophagous, saprophagous, xylomycetophagous, predators and commensals). We quantitatively assessed which properties of specialised networks were present in a complex networks involving different interacting types such as saproxylic community, and how they can be organised in trophic food webs. The architecture, interacting patterns and food web composition were evaluated along sub-networks, analysing their implications to network robustness from random and directed extinction simulations. A structure of large and cohesive modules with weakly connected nodes was observed throughout saproxylic sub-networks, composing the main food webs constituting this community. Insect-dependent sub-networks were more modular than wood-dependent sub-networks. Wood-dependent sub-networks presented higher species degree, connectance, links, linkage density, interaction strength, and were less specialised and more aggregated than insect-dependent sub-networks. These attributes defined high network robustness in wood-dependent sub-networks. Finally, our results emphasise the relevance of modularity, differences among interacting types and interrelations among them in modelling the structure of saproxylic communities and in determining their stability.

(Table A1) Annual biomass of plants and animals recorded on Bylot Island between 1993-2009

Determining the manner in which food webs will respond to environmental changes is difficult because the relative importance of top-down vs. bottom-up forces in controlling ecosystems is still debated. This is especially true in the Arctic tundra where, despite relatively simple food webs, it is still unclear which forces dominate in this ecosystem. Our primary goal was to assess the extent to which a tundra food web was dominated by plant-herbivore or predator--rey interactions. Based on a 17-year (1993-2009) study of terrestrial wildlife on Bylot Island, Nunavut, Canada, we developed trophic mass balance models to address this question. Snow Geese were the dominant herbivores in this ecosystem, followed by two sympatric lemming species (brown and collared lemmings). Arctic foxes, weasels, and several species of birds of prey were the dominant predators. Results of our trophic models encompassing 19 functional groups showed that <10% of the annual primary production was consumed by herbivores in most years despite the presence of a large Snow Goose colony, but that 20-100% of the annual herbivore production was consumed by predators. The impact of herbivores on vegetation has also weakened over time, probably due to an increase in primary production. The impact of predators was highest on lemmings, intermediate on passerines, and lowest on geese and shorebirds, but it varied with lemming abundance. Predation of collared lemmings exceeded production in most years and may explain why this species remained at low density. In contrast, the predation rate on brown lemmings varied with prey density and may have contributed to the high-amplitude, periodic fluctuations in the abundance of this species. Our analysis provided little evidence that herbivores are limited by primary production on Bylot Island. In contrast, we measured strong predator-prey interactions, which supports the hypothesis that this food web is primarily controlled by top-down forces. The presence of allochthonous resources subsidizing top predators and the absence of large herbivores may partly explain the predominant role of predation in this low-productivity ecosystem.

Behavior analysis of oithona in hypoxic/anoxic conditions

Senior thesis written for Oceanography 445

Using nitrogen stable isotope ratios ( d15N) of macroalgae to monitor changes in the extent of sewage impacts in Moreton Bay, Australia

During the past decade the use of stable isotopes to investigate transport pathways of nutrients in aquatic ecosystems has contributed new understanding and knowledge to many aspects of ecology; from the trophic structure of food webs to the spatial extent of nutrient discharges. At the same time aquatic monitoring programs around the world have become more interested in quantifying ecosystem health rather than simply measuring the physical and chemical properties of water (nutrients, pH, temperature and turbidity). A novel technique was initiated in 1998 as part of the development of the Ecosystem Health Monitoring Program in S.E. Queensland Australia (EHMP) using changes in the 15N value of the red macroalgae Catenella nipae, to indicate regions impacted by sewage nitrogen. Sewage plume mapping, using the 15N of C. nipae, has demonstrated that over the past 5 years there has been a large reduction in the magnitude and spatial extent of 15N enrichment at sites close to sewage treatment plants (STPs) discharging into Moreton Bay. This presentation will discuss how the 15N signatures of the C. nipae in the plume at the mouth of the Brisbane River have declined since it was first sampled in 1998 and will evaluate causes that may be responsible for these variations. A series of laboratory experiments were conducted to investigate how environmental conditions influence the 15N signature of C, nipae over the incubation period. These data will be used to discuss the observed in situ decline in 15N in an attempt to determine if the reduction can be attributed solely to improvements in the wastewater discharge.

Stable isotopes reveal complex changes in trophic relationships following nutrient addition in a coastal marine ecosystem

Complex links between the top-down and bottomup forces that structure communities can be disrupted by anthropogenic alterations of natural habitats.We used relative abundance and stable isotopes to examine changes in epifaunal food webs in seagrass (Thalassia testudinum) beds following 6 months of experimental nutrient addition at two sites in Florida Bay (USA) with different ambient fertility. At a eutrophic site, nutrient addition did not strongly affect food web structure, but at a nutrient-poor site, enrichment increased the abundances of crustacean epiphyte grazers, and the diets of these grazers became more varied. Benthic grazers did not change in abundance but shifted their diet away from green macroalgae + associated epiphytes and towards an opportunistic seagrass (Halodule wrightii) that occurred only in nutrient addition treatments. Benthic predators did not change in abundance, but their diets were more varied in enriched plots. Food chain length was short and unaffected by site or nutrient treatment, but increased food web complexity in enriched plots was suggested by increasingly mixed diets. Strong bottom-up modifications of food web structure in the nutrient-limited site and the limited top-down influences of grazers on seagrass epiphyte biomass suggest that, in this system, the bottom-up role of nutrient enrichment can have substantial impacts on community structure, trophic relationships, and, ultimately, the productivity values of the ecosystem.

Fatty acids and stable isotopes as indicators of early-life feeding and potential maternal resource dependency in the bull shark Carcharhinus leucas

The degree of reliance of newborn sharks on energy reserves from maternal resource allocation and the timescales over which these animals develop foraging skills are critical factors towards understanding the ecological role of top predators in marine ecosystems. We used muscle tissue stable carbon isotopic composition and fatty acid analysis of bull sharks Carcharhinus leucas to investigate early-life feeding ecology in conjunction with maternal resource dependency. Values of δ13C of some young-of-the-year sharks were highly enriched, reflecting inputs from the marine-based diet and foraging locations of their mothers. This group of sharks also contained high levels of the 20:3ω9 fatty acid, which accumulates during periods of essential fatty acid deficiency, suggesting inadequate or undeveloped foraging skills and possible reliance on maternal provisioning. A loss of maternal signal in δ13C values occurred at a length of approximately 100 cm, with muscle tissue δ13C values reflecting a transition from more freshwater/estuarine-based diets to marine-based diets with increasing length. Similarly, fatty acids from sharks >100 cm indicated no signs of essential fatty acid deficiency, implying adequate foraging. By combining stable carbon isotopes and fatty acids, our results provided important constraints on the timing of the loss of maternal isotopic signal and the development of foraging skills in relation to shark size and imply that molecular markers such as fatty acids are useful for the determination of maternal resource dependency.

A molecular and stable isotopic approach to investigate the importance of algal and detrital energy pathways in a freshwater marsh

The relative importance of algal and detrital energy pathways remains a central question in wetlands ecology. We used bulk stable isotope analysis and fatty acid composition to investigate the relative contributions of periphyton (algae) and floc (detritus) in a freshwater wetland with the goal of determining the inputs of these resource pools to lower trophic-level consumers. All animal samples revealed fatty acid markers indicative of both microbial (detrital) and algal origins, though the relative contributions varied among species. Vascular plant markers were in low abundance in most consumers. Detritivory is important for chironomids and amphipods, as demonstrated by the enhanced bacterial fatty acids present in both consumers, while algal resources, in the form of periphyton, likely support ephemeropteran larvae. Invertebrates such as amphipods and grass shrimp appear to be important resources for small omnivorous fish, while Poecilia latipinna appear to strongly use periphyton and Ephemeroptera larvae as food sources. Both P. latipinna and Lepomis spp. assimilated small amounts of vascular plant debris, possibly due to unintentional ingestion of floc while foraging for invertebrates and insect larvae. Physid snails, Haitia spp., were characterized by considerably different fatty acid compositions than other taxa examined, and likely play a unique role in Everglades’ food webs.

A synthesis of long-term research by the Florida Coastal Everglades LTER Program

This paper synthesizes research conducted during the first 5–6 years of the Florida Coastal Everglades Long-Term Ecological Research Program (FCE LTER). My objectives are to review our research to date, and to present a new central theme and conceptual approach for future research. Our research has focused on understanding how dissolved organic matter (DOM) from upstream oligotrophic marshes interacted with a marine source of the limiting nutrient, phosphorus (P), to control productivity in the oligohaline estuarine ecotone. We have been working along freshwater to marine transects in two drainage basins located in Everglades National Park (ENP). The Shark River Slough transect (SRS) has a direct connection to the Gulf of Mexico, providing this estuarine ecotone with a source of marine P. The oligohaline ecotone along our southern Everglades transect (TS/Ph), however, is separated from this marine P source by the Florida Bay estuary. We originally hypothesized an ecosystem productivity peak in the SRS ecotone, driven by the interaction of marine P and Everglades DOM, but no such productivity peak in the TS/Ph ecotone because of this lack of marine P. Our research to date has tended to show the opposite pattern, however, with many ecosystem components showing enhanced productivity in the TS/Ph ecotone, but not in the SRS ecotone. Water column P concentrations followed a similar pattern, with unexpectedly high P in the TS/Ph ecotone during the dry season. Our organic geochemical research has shown that Everglades DOM is more refractory than originally hypothesized. We have also begun to understand the importance of detrital organic matter production and transport to ecotone dynamics and as the base of aquatic food webs. Our future research will build on this substantial body of knowledge about these oligotrophic estuaries. We will direct our efforts more strongly on biophysical dynamics in the oligohaline ecotone regions. Specifically, we will be focusing on inputs to these regions from four primary water sources: freshwater Everglades runoff, net precipitation, marine inputs, and groundwater. We are hypothesizing that dry season groundwater inputs of P will be particularly important to TS/Ph ecotone dynamics because of longer water residence times in this area. Our organic geochemical, biogeochemical, and ecosystem energetics work will focus more strongly on the importance of detrital organics and will take advantage of a key Everglades Restoration project, scheduled for 2008 or 2009, that will increase freshwater inputs to our SRS transect only. Finally, we will also begin to investigate the human dimensions of restoration, and of a growing population in south Florida that will become increasingly dependent on the Everglades for critical ecosystem services (including fresh water) even as its growth presents challenges to Everglades sustainability.

Spatial and seasonal variability in elemental content, δ13C, and δ15N ofThalassia testudinum from South Florida and its implications for ecosystem studies

Elemental and isotopic composition of leaves of the seagrassThalassia testudinum was highly variable across the 10,000 km2 and 8 years of this study. The data reported herein expand the reported range in carbon:nitrogen (C:N) and carbon:phosphorus (C:P) ratios and δ13C and δ15N values reported for this species worldwide; 13.2–38.6 for C:N and 411–2,041 for C:P. The 981 determinations in this study generated a range of −13.5‰ to −5.2‰ for δ13C and −4.3‰ to 9.4‰ for δ15N. The elemental and isotope ratios displayed marked seasonality, and the seasonal patterns could be described with a simple sine wave model. C:N, C:P, δ13C, and δ15N values all had maxima in the summer and minima in the winter. Spatial patterns in the summer maxima of these quantities suggest there are large differences in the relative availability of N and P across the study area and that there are differences in the processing and the isotopic composition of C and N. This work calls into question the interpretation of studies about nutrient cycling and food webs in estuaries based on few samples collected at one time, since we document natural variability greater than the signal often used to imply changes in the structure or function of ecosystems. The data and patterns presented in this paper make it clear that there is no threshold δ15N value for marine plants that can be used as an unambiguous indicator of human sewage pollution without a thorough understanding of local temporal and spatial variability.

Carbon (d13c) and Nitrogen (d15n) Isotopic Discrimination in Mangroves in Florida Coastal Everglades as a Function of Environmental Stress

Isotope signatures of mangrove leaves can vary depending on discrimination associated with plant response to environmental stressors defined by gra­dients of resources (such as water and nutrient limitation) and regulators (such as salinity and sul­fide toxicity). We tested the variability of man­grove isotopic signatures (d13C and d15N) across a stress gradient in south Florida, using green leaves from four mangrove species collected at six sites. Mangroves across the landscape studied are stressed by resource and regulator gradients repre­sented by limited phosphorus concentrations com­bined with high sulfide concentrations, respec­tively. Foliar d13C ratios exhibited a range from ­ 24.6 to –32.7‰, and multiple regression analysis showed that 46% of the variability in mangrove d13C composition could be explained by the differ­ences in dissolved inorganic nitrogen, soluble reac­tive phosphorus, and sulfide porewater concentra­tions. 15N discrimination in mangrove species ranged from –0.1 to 7.7‰, and porewater N, salin­ity, and leaf N:Pa ratios accounted for 41% of this variability in mangrove leaves. The increase in soil P availability reduced 15N discrimination due to higher N demand. Scrub mangroves (<1.5 m tall) are more water-use efficient, as indicated by higher d13C; and have greater nutrient use efficiency ratios of P than do tall mangroves (5 to 10 m tall) existing in sites with greater soil P concentrations. The high variability of mangrove d13C and d15N across these resource and regulator gradients could be a con­founding factor obscuring the linkages between mangrove wetlands and estuarine food webs. These results support the hypothesis that landscape fac­tors may control mangrove structure and function, so that nutrient biogeochemistry and mangrove-based food webs in adjacent estuaries should ac­count for watershed-specific organic inputs.

Alligators and crocodiles as indicators for restoration of Everglades ecosystems

Alligators and crocodiles integrate biological impacts of hydrological operations, affecting them at all life stages through three key aspects of Everglades ecology: (1) food webs, (2) diversity and productivity, and (3) freshwater flow. Responses of crocodilians are directly related to suitability of environmental conditions and hydrologic change. Correlations between biological responses and environmental conditions contribute to an understanding of species’ status and trends over time. Positive or negative trends of crocodilian populations relative to hydrologic changes permit assessment of positive or negative trends in restoration. The crocodilian indicator uses monitoring parameters (performance measures) that have been shown to be both effective and efficient in tracking trends. The alligator component uses relative density (reported as an encounter rate), body condition, and occupancy rates of alligator holes; the crocodile component uses juvenile growth and hatchling survival. We hypothesize that these parameters are correlated with hydrologic conditions including depth, duration, timing, spatial extent and water quality. Salinity is a critical parameter in estuarine habitats. Assessments of parameters defined for crocodilian performance measures support these hypotheses. Alligators and crocodiles are the charismatic megafauna of the Everglades. They are both keystone and flagship species to which the public can relate. In addition, the parameters used to track trends are easy to understand. They provide answers to the following questions: How has the number of alligators or crocodiles changed? Are the animals fatter or thinner than they should be? Are the animals in the places (in terms of habitat and geography) where they should be? As surely as there is no other Everglades, no other single species defines the Everglades as does the American alligator. The Everglades is the only place in the world where both alligators and crocodiles exist. Crocodilians clearly respond to changes in hydrologic parameters of management interest. These relationships are easy to communicate and mean something to managers, decision makers, and the public. Having crocodilians on the list of system-wide, general indicators provides us with one of the most powerful tools we have to communicate progress of ecosystem restoration in Greater Everglades ecosystems to diverse audiences.

Organic Biogeochemistry of Detrital Flocculent Material (Floc) in a Subtropical, Coastal Wetland

Flocculent materials (floc), in aquatic systems usually consist of a non-consolidated layer of biogenic, detrital material relatively rich in organic matter which represents an important food-web component for invertebrates and fish. Thus, variations in its composition could impact food webs and change faunal structure. Transport, remineralization rates and deposition of floc may also be important factors in soil/sediment formation. In spite of its relevance and sensitivity to external factors, few chemical studies have been carried out on the biogeochemistry of floc material. In this study, we focused on the molecular characterization of the flocculent organic matter (OM), the assessment of its origin and its environmental fate at five stations along a freshwater to marine ecotone, namely the Taylor Slough, Everglades National Park (ENP), Florida. To tackle this issue, suspended, unconsolidated, detrital floc samples, soils/sediments and plants were analyzed for bulk properties, biomarkers and pigments. Both geochemical proxies and biomass-specific biomarkers were used to assess OM sources and transformations. Our results show that the detrital organic matter of the flocculent material is largely regulated by local vegetation inputs, ranging from periphyton, emergent and submerged plants and terrestrial plants such as mangroves, with molecular evidence of different degrees of diagenetic reworking, including fungal activity. Evidence is presented for both hydrodynamic transport of floc materials, and incorporation of floc OM into soils/sediments. However, some molecular parameters showed a decoupling between floc and underlying soil/sediment OM, suggesting that physical transport, incorporation and degradation/remineralization of OM in floc may be controlled by a combination of a variety of complex biogeochemical variables including hydrodynamic transport, hydroperiod characteristics, primary productivity, nutrient availability, and OM quality among others. Further investigations are needed to better understand the ecological role of floc in freshwater and coastal wetlands.

Comparison of snail density, standing stock, and body size between Caribbean karst wetlands and other freshwater ecosystems

Synthesizing data from multiple studies generates hypotheses about factors that affect the distribution and abundance of species among ecosystems. Snails are dominant herbivores in many freshwater ecosystems, but there is no comprehensive review of snail density, standing stock, or body size among freshwater ecosystems. We compile data on snail density and standing stock, estimate body size with their quotient, and discuss the major pattern that emerges. We report data from 215 freshwater ecosystems taken from 88 studies that we placed into nine categories. Sixty-five studies reported density, seven reported standing stock, and 16 reported both. Despite the breadth of studies, spatial and temporal sampling scales were limited. Researchers used 25 different sampling devices ranging in area from 0.0015 to 2.5 m2. Most ecosystem categories had similar snail densities, standing stocks, and body sizes suggesting snails shared a similar function among ecosystems. Caribbean karst wetlands were a striking exception with much lower density and standing stock, but large body size. Disparity in body size results from the presence of ampullariids in Caribbean karst wetlands suggesting that biogeography affects the distribution of taxa, and in this case size, among aquatic ecosystems. We propose that resource quality explains the disparity in density and standing stock between Caribbean karst wetlands and other categories. Periphyton in Caribbean karst wetlands has high carbon-to-phosphorous ratios and defensive characteristics that inhibit grazers. Unlike many freshwater ecosystems where snails are key grazers, we hypothesize that a microbial loop captures much of the primary production in Caribbean karst wetlands.

Does variation in movement tactics and trophic interactions among American alligators create habitat linkages?

1. Highly mobile top predators are hypothesized to spatially and/or temporally link disparate habitats through the combination of their movement and feeding patterns, but recent studies suggest that individual specialization in habitat use and feeding could keep habitats compartmentalized. 2.  We used passive acoustic telemetry and stable isotope analysis to investigate whether specialization in movement and feeding patterns of American alligators (Alligator mississippiensis) in an oligotrophic subtropical estuary created habitat linkages between marine and estuarine/freshwater food webs. 3.  Individual alligators adopted one of the three relatively distinct movement tactics that were linked to variation in diets. Fifty-six per cent of alligators regularly travelled from the upstream (freshwater/mid-estuary) areas into the downstream (marine-influenced) areas where salinities exceed those typically tolerated by alligators. Thirty-one per cent of the alligators made regular trips from the mid-estuarine habitat into the upstream habitat; 13% remained in the mid-estuary zone year-round. 4.  Stable isotopic analysis indicated that, unlike individuals remaining in the mid-estuary and upstream zones, alligators that used the downstream zone fed at least partially from marine food webs and likely moved to access higher prey abundance at the expense of salt stress. Therefore, ‘commuting’ alligators may link marine food webs with those of the estuary and marshes in the coastal Everglades and create an upstream vector for allochthonous nutrient inputs into the estuary. 5.  This study lends further support to the hypothesis that large-bodied highly mobile predators faced with trade-offs are likely to exhibit individual specialization leading to habitat linkages, rather than compartmentalization. However, the conditions under which this scenario occurs require further investigation.

Distribution of Diatoms and Development of Diatom-Based Models for Inferring Salinity and Nutrient Concentrations in Florida Bay and Adjacent Coastal Wetlands of South Florida (USA)

The composition and distribution of diatom algae inhabiting estuaries and coasts of the subtropical Americas are poorly documented, especially relative to the central role diatoms play in coastal food webs and to their potential utility as sentinels of environmental change in these threatened ecosystems. Here, we document the distribution of diatoms among the diverse habitat types and long environmental gradients represented by the shallow topographic relief of the South Florida, USA, coastline. A total of 592 species were encountered from 38 freshwater, mangrove, and marine locations in the Everglades wetland and Florida Bay during two seasonal collections, with the highest diversity occurring at sites of high salinity and low water column organic carbon concentration (WTOC). Freshwater, mangrove, and estuarine assemblages were compositionally distinct, but seasonal differences were only detected in mangrove and estuarine sites where solute concentration differed greatly between wet and dry seasons. Epiphytic, planktonic, and sediment assemblages were compositionally similar, implying a high degree of mixing along the shallow, tidal, and storm-prone coast. The relationships between diatom taxa and salinity, water total phosphorus (WTP), water total nitrogen (WTN), and WTOC concentrations were determined and incorporated into weighted averaging partial least squares regression models. Salinity was the most influential variable, resulting in a highly predictive model (r apparent 2  = 0.97, r jackknife 2  = 0.95) that can be used in the future to infer changes in coastal freshwater delivery or sea-level rise in South Florida and compositionally similar environments. Models predicting WTN (r apparent 2  = 0.75, r jackknife 2  = 0.46), WTP (r apparent 2  = 0.75, r jackknife 2  = 0.49), and WTOC (r apparent 2  = 0.79, r jackknife 2  = 0.57) were also strong, suggesting that diatoms can provide reliable inferences of changes in solute delivery to the coastal ecosystem.