Hydrology-mediated ecological function of a large wetland threatened by an invasive predator.

Invasive species are one of the greatest threats to ecosystems, disrupting ecosystem function and leading to the collapse and extinction of native species. While populations of native fishes in the Everglades are tied to the system's natural hydrological dynamics, Asian Swamp Eels (Monopterus albus/javanensis) are drought-resistant fish first reported from Florida in 1997 and the Everglades in 2007. Using a 26-year dataset that included a 13-year baseline period prior to swamp eel arrival in Taylor Slough, we assessed population changes of common small fishes and decapods that are important prey for larger vertebrate predators. After invasion, populations of two crayfishes collapsed by >95 %, two fishes declined by >80 %, two fishes had intermediate declines of 44-66 %, and three species remained unchanged. Species most strongly reduced were those dependent on predator-free habitats at the onset of the wet season, indicating drought-resistant swamp eels have introduced novel predator effects and disrupted the hydrology-mediated production of aquatic animals that are prey for many larger predators. Ongoing Everglades restoration is designed to restore hydrological conditions that support production of crayfishes and fishes, and nesting wading birds reliant on them. Water management may have facilitated the invasion of swamp eels. Our results suggest that the continued spread of swamp eels may result in adverse consequences for Everglades trophic dynamics and potentially diminish benefits expected from the $20B+ restoration.

A skewed literature: Few studies evaluate the contribution of predation-risk effects to natural field patterns.

A narrative in ecology is that prey modify traits to reduce predation risk, and the trait modification has costs large enough to cause ensuing demographic, trophic and ecosystem consequences, with implications for conservation, management and agriculture. But ecology has a long history of emphasising that quantifying the importance of an ecological process ultimately requires evidence linking a process to unmanipulated field patterns. We suspected that such process-linked-to-pattern (PLP) studies were poorly represented in the predation risk literature, which conflicts with the confidence often given to the importance of risk effects. We reviewed 29 years of the ecological literature which revealed that there are well over 4000 articles on risk effects. Of those, 349 studies examined risk effects on prey fitness measures or abundance (i.e., non-consumptive effects) of which only 26 were PLP studies, while 275 studies examined effects on other interacting species (i.e., trait-mediated indirect effects) of which only 35 were PLP studies. PLP studies were narrowly focused taxonomically and included only three that examined unmanipulated patterns of prey abundance. Before concluding a widespread and influential role of predation-risk effects, more attention must be given to linking the process of risk effects to unmanipulated patterns observed across diverse ecosystems.

Phylogeographic reconstruction of the marbled crayfish origin.

The marbled crayfish (Procambarus virginalis) is a triploid and parthenogenetic freshwater crayfish species that has colonized diverse habitats around the world. Previous studies suggested that the clonal marbled crayfish population descended as recently as 25 years ago from a single specimen of P. fallax, the sexually reproducing parent species. However, the genetic, phylogeographic, and mechanistic origins of the species have remained enigmatic. We have now constructed a new genome assembly for P. virginalis to support a detailed phylogeographic analysis of the diploid parent species, Procambarus fallax. Our results strongly suggest that both parental haplotypes of P. virginalis were inherited from the Everglades subpopulation of P. fallax. Comprehensive whole-genome sequencing also detected triploid specimens in the same subpopulation, which either represent evolutionarily important intermediate genotypes or independent parthenogenetic lineages arising among the sexual parent population. Our findings thus clarify the geographic origin of the marbled crayfish and identify potential mechanisms of parthenogenetic speciation.

Morphological characterization and genotyping of the marbled crayfish and new evidence on its origin.

The obligately parthenogenetic marbled crayfish, Procambarus virginalis, is the first formally described asexual species of the Crustacea Decapoda. It is a triploid descendant of the sexually reproducing slough crayfish, Procambarus fallax. Here we describe the morphology of cultured and wild marbled crayfish of wide size ranges in detail and photodocument all taxonomically relevant characters. Some morphological traits and coloration showed considerable variation within populations despite the monoclonal nature of marbled crayfish. There were also significant differences between wild and laboratory populations with respect to body proportions, coloration and spination. Comparison with Procambarus fallax revealed no qualitative morphological characters that unambiguously identify the marbled crayfish. Analysis of the mitochondrial cytochrome c oxidase subunit I gene (COI) and nuclear microsatellites of marbled crayfish and Procambarus fallax from different sources indicated that the tri-allelic microsatellite PclG-02 is better suitable than COI to identify the marbled crayfish. A respective identification key is provided. The COI and microsatellites of Procambarus fallax from different areas of Florida and southern Georgia suggest that the parents of the first marbled crayfish may have come from northern Union County, northern Florida.

Correction to: System productivity alters predator sorting of a size-structured mixed prey community.

There was an error in the abstract of the original publication. The 9th sentence of abstract should be:"Our results are inconsistent with predictions about the sorting effects of predators across productivity gradients because the more resistant prey dominated at low productivity."

System productivity alters predator sorting of a size-structured mixed prey community.

Predator-prey interactions are often size-structured and focused on smaller vulnerable size classes. Predators are also predicted to sort prey communities according to relative vulnerabilities. Increased system productivity and juvenile growth may benefit some species more than others, making relative vulnerability non-static and growth-mediated. We hypothesized that increased system productivity would weaken juvenile-stage predation generally, and potentially shift the community sorting effects of a predator. Using replicated wetland mesocosms we quantified the effects of a generalist size-specific crayfish predator (Procambarus fallax) on juveniles of two species of apple snails (Pomacea spp.) under two levels of system productivity (low vs. high). After 6 weeks of exposure, we quantified predator and productivity effects on snail survival, biomass, and composition of the assemblage. Crayfish depressed the final density and biomass of snails, and sorted the assemblage, selectively favoring survival of the native P. paludosa over the intrinsically more vulnerable invasive P. maculata. Both snails grew faster at higher productivity, but growth differentially increased survival of the invasive snail in the presence of crayfish and weakened the sorting effect. The native P. paludosa hatches at a larger less vulnerable size than the invasive P. maculata, but higher productivity reduced the relative advantage of P. paludosa. Our results are inconsistent with predictions about the sorting effects of predators across productivity gradients because the more resistant prey dominated at low productivity. Our findings highlight that the relative vulnerabilities of prey to a common predator are not always fixed, but can be growth-mediated.

Hydrological disturbance diminishes predator control in wetlands.

Effects of predators on prey populations can be especially strong in aquatic ecosystems, but disturbances may mediate the strength of predator limitation and even allow outbreaks of some prey populations. In a two-year study we investigated the numerical responses of crayfish (Procambarus fallax) and small fishes (Poeciliidae and Fundulidae) to a brief hydrological disturbance in replicated freshwater wetlands with an experimental drying and large predatory fish reduction. The experiment and an in situ predation assay tested the component of the consumer stress model positing that disturbances release prey from predator limitation. In the disturbed wetlands, abundances of large predatory fish were seasonally reduced, similar to dynamics in the Everglades (southern Florida). Densities of small fish were unaffected by the disturbance, but crayfish densities, which were similar across all wetlands before drying, increased almost threefold in the year after the disturbance. Upon re-flooding, juvenile crayfish survival was inversely related to the abundance of large fish across wetlands, but we found no evidence for enhanced algal food quality. At a larger landscape scale (500 km2 of the Everglades), crayfish densities over eight years were positively correlated with the severity of local dry disturbances (up to 99 days dry) during the preceding dry season. In contrast, densities of small-bodied fishes in the same wetlands were seasonally depressed by dry disturbances. The results from our experimental wetland drought and the observations of crayfish densities in the Everglades represent a large-scale example of prey population release following a hydrological disturbance in a freshwater ecosystem. The conditions producing crayfish pulses in the Everglades appear consistent with the mechanics of the consumer stress model, and we suggest crayfish pulses may influence the number of nesting wading birds in the Everglades.

Consumptive effects of fish reduce wetland crayfish recruitment and drive species turnover.

Predators and dry-disturbances have pronounced effects on invertebrate communities and can deterministically affect compositional turnover between discrete aquatic habitats. We examined the effect of sunfish (Lepomis spp.) predators on two native crayfish, Procambarus alleni and P. fallax, that regionally coexist in an expansive connected wetland in order to test the hypotheses that sunfish predation limits crayfish recruitment (both species) and that it disproportionately affects P. alleni, the species inhabiting temporary wetlands. In replicate vegetated wetlands (18.6 m(2)), we quantified summertime crayfish recruitment and species composition across an experimental gradient of sunfish density. Separately, we quantified effects of sunfish on crayfish growth, conducted a complimentary predation assay in mesocosms, and compared behavior of the two crayfish. Sunfish reduced P. alleni summertime recruitment by >99% over the full sunfish gradient, and most of the effect was caused by low densities of sunfish (0.22-0.43 m(-2)). P. alleni dominated wetlands with few or no sunfish, but the composition shifted towards P. fallax dominance in wetlands with abundant sunfish. P. fallax survived better than P. alleni over 40 h in smaller mesocosms stocked with warmouth. Sunfish reduced P. fallax recruitment 62% in a second wetland experiment, but growth rates of caged crayfish (both species) were unaffected by sunfish presence, suggesting predatory effects were primarily consumptive. Consistent with life histories of relatively fish-sensitive invertebrates, P. alleni engaged in more risky behaviors in the laboratory. Our results indicate that sunfish predators limit recruitment of both species, but disproportionately remove the more active and competitively dominant P. alleni. Spatially and temporally variable dry-disturbances negatively co-varying with sunfish populations allow for regional coexistence of these two crayfish and may release populations of either species from control by predatory fishes.

The role of omnivorous crayfish in littoral communities.

Large omnivorous predators may play particularly important roles determining the structure of communities because of their broad diets and simultaneous effects on multiple trophic levels. From June 2001 to June 2002 we quantified community structure and ecosystem attributes of six newly establishing freshwater ponds (660 m2 each) after populations of omnivorous crayfish (Orconectes virilis) were introduced to three of the ponds. Crayfish preyed heavily on fish eggs in this experiment, which reduced recruitment of young-of-year fish. This effect indirectly enhanced zooplankton biomass in crayfish ponds. Phytoplankton abundance exhibited a more complex pattern and was probably influenced by non-trophic (e.g., bioturbation) effects of crayfish. Peak dissolved oxygen levels were lower in the crayfish ponds indicating that they had lower primary production: respiration ratios. Metaphytic algae were strongly affected by crayfish presence; filamentous greens quickly disappeared and the blue-green Gleotrichia (a less preferred food item) eventually dominated the composition in crayfish ponds. Chara vulgaris and vascular macrophytes established 34% cover in control ponds by June 2002, but were not able to establish in crayfish ponds. Two important periphyton herbivores (tadpoles and gastropods) were absent or significantly reduced in the crayfish ponds, but periphyton differences were temporally variable and not easily explained by a simple trophic cascade (i.e., crayfish-snails and tadpoles-periphyton). Our results indicate that crayfish can have dramatic direct and indirect impacts on littoral pond communities via feeding links with multiple trophic levels (i.e., fish, invertebrates, and plants) and non-trophic activities (bioturbation). Although the effects of omnivorous crayfish on littoral communities can be large, their complex effects do not fit neatly into current theories about trophic interactions or freshwater community structure.

Feeding preferences and performance of an aquatic lepidopteran on macrophytes: plant hosts as food and habitat.

Although host preferences in phytophagous insects may be generated by several factors, few studies have simultaneously examined several potential host choice determinants. In this study we tested the impact of the following potential host choice determinants on host preference of the semi-aquatic lepidopteran Munroessa gyralis (Pyralidae): growth on different host plants; protein content, polyphenolic content, toughness, and chemical extracts of different host plants; prior feeding experience; and predation pressure on the caterpillar by fishes. Two water lilies, Brasenia schreberi and Nymphaea odorata, were preferred in cafeteria-style feeding experiments over 14 other species of vascular plants. The most preferred water lily (Brasenia) also afforded the fastest growth relative to three other species on which growth was measured. Feeding preferences across species were unrelated to protein content, polyphenolic content, or toughness. Domiciles constructed by caterpillars from leaf fragments were protective from field assemblages of fishes, but domiciles made from preferred or unpreferred host species conferred no significant protection from fish in the laboratory. Caterpillars responded positively to chemical cues of water lilies, and prior feeding experience increased preference for an otherwise unpreferred water lily (Nuphar advena) within the life-span of individual caterpillars. M. gyralis is a generalist herbivore exhibiting modest preference induction and preferences for and among members of the family Nymphaeaceae. Our results suggest that relative growth rates, chemical cues, and previous feeding experience are important factors determining feeding preference. Protein content, polyphenolic content, and toughness appear less important, and the importance of fish predators remains in question. As pupation seems to occur exclusively on Nymphaea, we suggest that host use may be restricted due to life-stage-specific developmental constraints that are not apparent from the results of growth or preference assays. It is currently unknown how often specific life-stages may restrict host use, but our work suggests this as a potentially important area of inquiry.