Prevalence of malformed frogs in Kaoping and Tungkang river basins of southern Taiwan

In this study we found many amphibians with bizarre appearances, known as malformations in Pingtung County southern Taiwan. For this investigation we collected frogs inhabiting the Kaoping and Tungkang river watersheds between February 2006 and June 2007. Among the total number of 10,909 normal frogs (i.e., anurans) collected during the investigation period, the Indian rice frogs (Rana limnocharis) account for the greatest number next is the Chinese bullfrog (Rana rugulosa). Of all the 244 captured malformed frogs, the Indian rice frog account for the greatest proportion. These malformed frogs have their main distribution in upstream areas of these two rivers. Our result indicates that the appearance rate of malformed frogs is 1.8% in the upstream reaches of the Kaoping River and 2.6%, and 0.8%, respectively in the upstream and midstream reaches of the Tungkang river. The most-commonly-found malformation is the lack of palms, followed by the lack of appendages, exostosis, and a malformed appendicular. It is, therefore, reasonable to speculate that the causes for the malformation may be related to the increased organic pollutants and agricultural chemicals used in the upstream reaches of these two rivers.     

Alterations of biochemical parameters in malformed Indian rice frogs, Rana limnocharis from Southern Taiwan

The purpose of this study is to investigate the factors that cause malformed frogs in upstream Kaoping river (KP site) and Tungkang river (T site) of Southern Taiwan. In this experiment, the activities of monooxygenase (MO), glutathione-S-transferase (GST), acetylcholinesterase (AchE) as well as the concentration of vitellogenin (Vg) in the liver were measured. Results show that activities of MO, GST and AchE, and Vg levels in normal frogs (male/female) were 0.09 +/- 0.02/0.09+/-0.01 deltaA min(-1) mg(-1) protein, 0.12 +/- 0.04/0.13 +/- 0.04 deltaA min(-1) mg(-1) protein, 6.13 +/- 2.69/6.01 +/- 2.09 U mg(-1) protein and 0.87 +/- 0.42/2.18 +/- 0.50 microg mg(-1) protein, respectively. Activities of MO, GST and AchE, and Vg levels in malformed frogs (male/female) were 0.15 +/- 0.04/0.21 +/- 0.07 deltaA min(-1) mg(-1) protein, 0.27 +/- 0.08/0.30 +/- 0.12 deltaA min(-1) mg(-1) protein, 4.59 +/- 2.71/5.19 +/- 3.74 U mg(-1) protein and 1.46 +/- 0.61/3.15 +/- 0.88 microg mg(-1) protein, respectively in KP site, and were 0.16 +/- 0.69/0.1 +/- 80.07 deltaA min(-1) mg(-1) protein, 0.21 +/- 0.07/0.24 +/- 0.08 deltaA min(-1) mg(-1) protein, 5.13 +/- 4.58/3.94 +/- 1.33 U mg(-1) protein and 2.23 +/- 1.47/4.11 +/- 1.63 microg mg(-1) protein, respectively in T site. These results indicate that male and female malformed frogs in both rivers upstream are found with higher activities. No significant difference in AchE activity was found between normal and malformed frogs in this investigation. It is therefore reasonable to speculate that the organic chemicals released from agricultural activities are presumable the main factors that lead to the malformation of frogs.     

Use of polyvinyl chloride pipes (PVC) as potential artificial shelters for amphibians in a coastal plain forest of southeastern Brazil

We studied the use of polyvinil chloride pipes (PVC pipes) by frogs as artificial shelters in a Coastal Plain forest in the Vale Natural Reserve, north of Espírito Santo state, southeastern Brazil from June 2015 to May 2016. We used 14 plots distributed in the Coastal Plain Forest following the RAPELD methodology of sampling. The frogs were sampled in transects along the plots, using the active search method, with visual and auditory sampling. We recorded two species of frogs (Aparasphenodon brunoi and Scinax argyreornatus) using PVC pipes as shelters in the plots in the Forest. Of the 468 PVC pipes established along plots, 38 of them (8.1%) were used as shelter by the frogs. For both frog species, about 33% of microhabitats used in the forest were the interior of PVC pipes. We concluded that the PVC pipes might constitute artificial shelters for at least two of the frog species living in the community of the Coastal Plain Forest, potentially being of relevance in conservation programs. In altered areas and those in which there has been the loss of types of microhabitats, the addition of these PVC pipes as artificial shelters may represent additional and complementary microhabitats to keep or increase the population stocks of arboreal and semi-arboreal frog species and having a size that allow them entering the shelter. These artificial shelters can play an important role in conservation of these frogs in fragmented areas where the availability of the microhabitats of these frog species were locally reduced.     

Inf luence of Season and a Sympatric Congener on Habitat Use by Stephens' Kangaroo Rat

We examined habitat use by the endangered Stephens' kangaroo rat ( Dipodomys stephensi) in different seasons and assessed whether this may be influenced by a sympatric congener, the Pacific kangaroo rat (Dipodomys agilis). Trapping on three plots over 2 years revealed these species were rarely captured at the same trap stations. Spatial segregation was highly significant when both species were at high density. The spatial distribution of these species was temporally stable where both species were relatively abundant, but where D. agilis was relatively uncommon the distribution of D. stephensi varied from one census to another. The abundance of three microhabitats (grass, debris, and bare ground) followed a regular seasonal pattern of variation that was consistent across 2 years of substantially different rainfall. A canonical discriminant analysis showed that the five quantified microhabitats (those above and bush and rock cover) provided highly significant discrimination between the trap stations occupied by the two species. Dipodomys stephensi was associated with trap stations where grass cover and bare ground were abundant but where bush and rock were uncommon. Dipodomys agilis was associated with stations that had large amounts of bare ground and average abundances of bush and rock cover. The spatial segregation of these species appears to be mediated by habitat preferences; D. stephensi prefers grassland and D. agilis prefers sage scrub. This suggests that habitat management for D. stephensi should include (1) controlling the spread of shrubs into grassland and (2) creating dispersal corridors of open habitat to link areas of suitable habitat where none presently exist. Each of these options may be needed to maintain viable populations in all reserves designated for the conservation of D. stephensi.     

Effects of Introduced Salmonids on a Montane Population of Iberian Frogs

Abstract:  Amphibians are declining worldwide because of multiple factors, including human-mediated introduction of fishes into naturally fishless areas. Although several studies have focused on the effect of exotic fishes on native amphibians breeding in ponds or lakes, little is known about their effects on stream-breeding species. We studied the effects of introductions of native brown trout ( Salmo trutta ) and exotic brook trout ( Salvelinus fontinalis ) on the stream-breeding, endemic Iberian frog (  Rana iberica ) in a protected area in central Spain. We assessed occurrence patterns of tadpoles and salmonids and compared habitat use of the three species. We also determined experimentally whether chemical cues from salmonids elicited antipredator behavior in tadpoles. Finally, we assessed the relative influence of tadpole habitat preferences, differences in salmonid species, and invasion geography on tadpole occurrence. Despite widely overlapping habitat preferences, tadpoles and trout did not coexist, with the former restricted to fishless habitats. Tadpoles detected chemical cues from both trout species and reacted by decreasing their activity, although the response toward the native brown trout was stronger. The residual distribution of Iberian frogs in Peñalara is better explained by the geography of fish invasions than by the fish species involved. Measures such as fish extirpation from certain areas, aimed at recovering lost habitat and improving connectivity among remaining populations of Iberian frogs, seem critical for the species' long-term survival in central Spain.     

Terrestrial habitat selection and strong density-dependent mortality in recently metamorphosed amphibians

To predict the effects of terrestrial habitat change on amphibian populations, we need to know how amphibians respond to habitat heterogeneity, and whether habitat choice remains consistent throughout the life-history cycle. We conducted four experiments to evaluate how the spatial distribution of juvenile wood frogs, Rana sylvatica (including both overall abundance and localized density), was influenced by habitat choice and habitat structure, and how this relationship changed with spatial scale and behavioral phase. The four experiments included (1) habitat manipulation on replicated 10-ha landscapes surrounding breeding pools; (2) short-term experiments with individual frogs emigrating through a manipulated landscape of 1 m wide hexagonal patches; and habitat manipulations in (3) small (4-m2); and (4) large (100-m2) enclosures with multiple individuals to compare behavior both during and following emigration. The spatial distribution of juvenile wood frogs following emigration resulted from differences in the scale at which juvenile amphibians responded to habitat heterogeneity during active vs. settled behavioral phases. During emigration, juvenile wood frogs responded to coarse-scale variation in habitat (selection between 2.2-ha forest treatments) but not to fine-scale variation. After settling, however, animals showed habitat selection at much smaller scales (2-4 m2). This resulted in high densities of animals in small patches of suitable habitat where they experienced rapid mortality. No evidence of density-dependent habitat selection was seen, with juveniles typically choosing to remain at extremely high densities in high-quality habitat, rather than occupying low-quality habitat. These experiments demonstrate how prediction of the terrestrial distribution of juvenile amphibians requires understanding of the complex behavioral responses to habitat heterogeneity. Understanding these patterns is important, given that human alterations to amphibian habitats may generate extremely high densities of animals, resulting in high density-dependent mortality.     

Parasites as conservation tools

Parasite success typically depends on a close relationship with one or more hosts; therefore, attributes of parasitic infection have the potential to provide indirect details of host natural history and are biologically relevant to animal conservation. Characterization of parasite infections has been useful in delineating host populations and has served as a proxy for assessment of environmental quality. In other cases, the utility of parasites is just being explored, for example, as indicators of host connectivity. Innovative studies of parasite biology can provide information to manage major conservation threats by using parasite assemblage, prevalence, or genetic data to provide insights into the host. Overexploitation, habitat loss and fragmentation, invasive species, and climate change are major threats to animal conservation, and all of these can be informed by parasites.     

Parasite distribution and virulence: implications for parasite-mediated sexual selection

The Hamilton-Zuk hypothesis states that females choosing males with more developed secondary sexual traits, i.e. brighter males, achieve greater fitness if variability in brightness reflects heritable variation in resistance to parasites. However, several factors will affect the likelihood that parasites play a role in sexual selection in given species. Here, using simple models, we show that because of parasite aggregation on a few hosts, only few breeding males would suffer from reductions in brightness due to parasites. Only in cases where parasites are abundant and show low levels of aggregation among their hosts would there be sufficient variability in brightness among breeding males for female choice of bright, resistant males to evolve. In addition, sufficient parasite-induced variability in brightness among breeding males will only occur in host-parasite systems where pathology is linearly related to the number of parasites per host. The presence of males that are uninfected and bright but genetically susceptible to parasites will also influence the fitness advantages obtained by females choosing bright males. If genetic immunity against parasites is rare in the host population, females can probably only benefit from choosing bright males if parasites are common and little aggregated among males. These results greatly limit the generality of the Hamilton-Zuk hypothesis, and suggest that only a small fraction of host-parasite associations could promote the evolution of host mate choice for resistance based on brightness. Correspondence to: R. Poulin     

Common frogs response to agrochemicals contamination in coffee plantations,Western Ghats,India

This study highlights the health status of frogs in coffee plantations of Western Ghats. Coffee is one of the major commercial crops in the Western Ghats of India. In these plantations, farmers extensively use agrochemicals including pesticides and synthetic fertilisers. The practice of agrochemical usage can negatively affect non-target organisms like amphibians. The area has reported many amphibian species living in and around plantations. A total of 334 frogs from reference sites (10 abnormal frogs) and a total of 258 frogs (19 abnormal frogs) were collected from contaminated sites. We considered major biomarkers like hepato-somatic index, gonado-somatic index, morphological abnormalities and acetylcholinestarse activities for evaluating the health status of frogs. In our study, the hepato-somatic index and gonado-somatic index were found to be higher in a grochemical contaminated sites. Whereas, acetycholine esterase activity in brain and liver tissues of frogs in contaminated sites were low compared to reference sites. High rate of incidence of morphological abnormality (7.36%), increased hepato-somatic index (14.86%) and gonado-somatic index (male; 8.88% and female; 17.51%) and low acetylcholine esterase activities in brain (41.5%) and liver (46.9%) indicates the lower health status of frogs living in coffee plantations that are regularly treated with agrochemicals.     

Shifted balance of risk and cost after autotomy affects use of cover,escape, activity,and foraging in the keeled earless lizard (<Emphasis Type="Italic">Holbrookia propinqua</Emphasis>)

Autotomy of expendable body parts is often a successful last-chance antipredatory defense for a variety of invertebrates and vertebrates, especially lizards. However, loss of a body part imposes costs and risks, some of which may be reduced by modifying subsequent behavior. I studied effects of experimentally induced autotomy of the tail in the keeled earless lizard (Holbrookia propinqua) on use of cover, escape behavior, activity, and feeding. Autotomized lizards stayed closer to plant cover than sham-autotomized lizards (having intact tails) when approached by an investigator simulating a predator. Autotomized males, but not females, fled further than sham-autotomized lizards. Autotomy did not affect the distance from an approaching predator when the lizards began to flee. After autotomy, the number of days on which lizards were active differed between sexes and lizards made fewer attempts to catch prey per unit time than lizards with intact tails. Tail loss did not affect the percentage of time spent moving. Staying closer to refuge, fleeing further after autotomy, and decreasing surface activity may compensate for decreased maximum escape speed or agility, and may reflect use of a greater margin of safety when the tail is unavailable for autotomy. Decrease in surface activity may be comparable to use of safer microhabitats reported for other lizards and damselflies subsequent to autotomy. Several possible reasons for the lower rate of feeding attempts by autotomized lizards are discussed.     

Temporal, spatial, and between-host comparisons of patterns of parasitism in lake zooplankton

In nature, multiple parasite species infect multiple host species and are influenced by processes operating across different spatial and temporal scales. Data sets incorporating these complexities offer exciting opportunities to examine factors that shape epidemics. We present a method using generalized linear mixed models in a multilevel modeling framework to analyze patterns of variances and correlations in binomially distributed prevalence data. We then apply it to a multi-lake, multiyear data set involving two Daphnia host species and nine microparasite species. We found that the largest source of variation in parasite prevalence was the species identities of host-parasite pairs, indicating strong host-parasite specificity. Within host-parasite combinations, spatial variation (among lakes) exceeded interannual variation. This suggests that factors promoting differences among lakes (e.g., habitat characteristics and species interactions) better explain variation in peak infection prevalence in our data set than factors driving differences among years (e.g., climate). Prevalences of parasites in D. dentifera were more positively correlated than those for D. pulicaria, suggesting that similar factors influenced epidemic size among parasites in D. dentifera. Overall, this study demonstrates a method for parsing patterns of variation and covariation in infection prevalence data, providing greater insight into the relative importance of different underlying drivers of parasitism.     

The Contribution of Habitat Loss to Changes in Body Size, Allometry, and Bilateral Asymmetry in Two Eleutherodactylus Frogs from Puerto Rico

Abstract:  Amphibian populations have been declining worldwide and the exact mechanisms underlying these changes are not well understood. We examined environmentally induced phenotypic changes that may reflect ongoing stresses on individuals and therefore their ability to persist in increasingly changing landscapes. Specifically, we evaluated the contribution of habitat loss on the size, allometry, and levels of fluctuating asymmetry of Eleutherodactylus antillensis and E. coqui , 2 common species that are endemic to Puerto Rico. We x-rayed frogs collected at 9 sites that differed in the amount of forest cover and measured their snout-vent, radio-ulna, femur, and tibio-fibula lengths. E. antillensis and E. coqui were smaller in the highly disturbed (≤20% forest cover) than in the intermediately (20–70% forest cover) and little-disturbed (≥70% forest cover) landscapes. In E. antillensis but not in E. coqui , the slope and intercept of the curves relating snout-vent length with the length of the 3 bones differed with degree of forest cover, suggesting an effect of habitat loss on body shape. In E. antillensis and E. coqui , differences between right and left sides corresponded to true fluctuating asymmetry; however, only the radio-ulna length of E. coqui showed a trend toward an increase in fluctuating asymmetry with habitat loss. Because body size scales with a variety of physiological, life history, and ecological traits, conservation programs aimed at monitoring morphological changes in amphibians may help in understanding the mechanisms that contribute to their persistence in changing environments.     

Genetic structure among coastal tailed frog populations at Mount St. Helens is moderated by post-disturbance management

Catastrophic disturbances often provide "natural laboratories" that allow for greater understanding of ecological processes and response of natural populations. The 1980 eruption of the Mount St. Helens volcano in Washington, USA, provided a unique opportunity to test biotic effects of a large-scale stochastic disturbance, as well as the influence of post-disturbance management. Despite severe alteration of nearly 600 km2 of habitat, coastal tailed frogs (Ascaphus truei) were found within a portion of the blast area five years after eruption. We investigated the genetic source of recolonization within the blast area and tested whether post-eruption salvage logging and subsequent tree planting influenced tailed frog movement patterns. Our results support widespread recolonization across the blast area from multiple sources, as all sites are grouped into one genetic cluster. Landscape genetic models suggest that gene flow through the unmanaged portion of the blast area is influenced only by distance between sites and the frost-free period (r2 = 0.74). In contrast, gene flow pathways within the blast area where salvage logging and replanting occurred post-eruption are strongly limited (r2 = 0.83) by the physiologically important variables of heat load and precipitation. These data suggest that the lack of understory and coarse wood (downed and standing dead tree boles) refugia in salvaged areas may leave frogs more susceptible to desiccation and mortality than those frogs moving through the naturally regenerated area. Simulated populations based on the landscape genetic models show an increase in the inbreeding coefficient in the managed area relative to the unmanaged blast area. In sum, we show surprising resilience of an amphibian species to a catastrophic disturbance, and we suggest that, at least for this species, naturally regenerating habitat may better maintain long-term genetic diversity of populations than actively managed habitat.     

Genetic relatedness and eusociality: parasite-mediated selection on the genetic composition of groups

Summary Contrary to the expectations of kin selection theory, intracolony relatedness in eusocial insects is often low. We examined the idea that associations of low relatedness (high genetic variability) may be advantageous because of negative frequency-dependent selection on common host phenotypes by rapidly evolving parasites and pathogens. Using the natural host-parasite system of the bumble bee Bombus terrestris and its intestinal trypanosome Crithidia bombi, we investigated the transmission properties of parasites in host groups. Within naturally infested nests and in artificially constructed groups of workers, prevalence of infestation increased with time of exposure (Table 1). The susceptibility of isolated groups of workers to the parasites to which they were exposed differed with identity and natural infestation of their nest of origin (Table 2). In addition, those workers that were related to the individual introducing an infection to their group were more likely to become infested than were unrelated workers (Table 3). Although the bumble bee workers in experimental boxes appeared to differ in behavior toward kin and non-kin, making more physical contacts with kin, we found no discernible relationship between number of physical contacts and prevalence of infestation in a group. Therefore, we conclude that differences in parasite transmission reflected interactions among different host and parasite phenotypes. This system thus demonstrates the factors necessary for negative frequency-dependent selection by parasites on common host phenotypes - variability for susceptibility and infectiousness in host and parasite populations and similarity for these traits among related individuals. If, as we show here, high genetic relatedness within groups enhances parasite transmission, kin directed altruism may increase the risk of contracting parasites and infectious diseases. Therefore, parasites and pathogens may be an important force moderating the genetic structure of social groups. Offprint requests to: J.A. Shykoff at the present address     

Effects of Landscape Composition and Wetland Fragmentation on Frog and Toad Abundance and Species Richness in Iowa and Wisconsin, U.S.A.

Abstract: Management of amphibian populations to reverse recent declines will require defining high-quality habitat for individual species or groups of species, followed by efforts to retain or restore these habitats on the landscape. We examined landscape-level habitat relationships for frogs and toads by measuring associations between relative abundance and species richness based on survey data derived from anuran calls and features of land-cover maps for Iowa and Wisconsin. The most consistent result across all anuran guilds was a negative association with the presence of urban land. Upland and wetland forests and emergent wetlands tended to be positively associated with anurans. Landscape metrics that represent edges and patch diversity also had generally positive associations, indicating that anurans benefit from a complex of habitats that include wetlands. In Iowa the most significant associations with relative abundance were the length of the edge between wetland and forest ( positive) and the presence of urban land (negative). In Wisconsin the two most significant associations with relative abundance were forest area and agricultural area ( both positive). Anurans had positive associations with agriculture in Wisconsin but not in Iowa. Remnant forest patches in agricultural landscapes may be providing refuges for some anuran species. Differences in anuran associations with deep water and permanent wetlands between the two states suggest opportunities for management action. Large-scale maps can contribute to predictive models of amphibian habitat use, but water quality and vegetation information collected from individual wetlands will likely be needed to strengthen those predictions. Landscape habitat analyses provide a framework for future experimental and intensive research on specific factors affecting the health of anurans.     

Effects of flow regimes altered by dams on survival, population declines, and range-wide losses of California river-breeding frogs

Widespread alteration of natural hydrologic patterns by large dams combined with peak demands for power and water delivery during summer months have resulted in frequent aseasonal flow pulses in rivers of western North America. Native species in these ecosystems have evolved with predictable annual flood-drought cycles; thus, their likelihood of persistence may decrease in response to disruption of the seasonal synchrony between stable low-flow conditions and reproduction. We evaluated whether altered flow regimes affected 2 native frogs in California and Oregon (U.S.A.) at 4 spatial and temporal extents. We examined changes in species distribution over approximately 50 years, current population density in 11 regulated and 16 unregulated rivers, temporal trends in abundance among populations occupying rivers with different hydrologic histories, and within-year patterns of survival relative to seasonal hydrology. The foothill yellow-legged frog (Rana boylii), which breeds only in flowing water, is more likely to be absent downstream of large dams than in free-flowing rivers, and breeding populations are on average 5 times smaller in regulated rivers than in unregulated rivers. Time series data (range = 8 - 19 years) from 5 populations of yellow-legged frogs and 2 populations of California red-legged frogs (R. draytonii) across a gradient of natural to highly artificial timing and magnitude of flooding indicate that variability of flows in spring and summer is strongly correlated with high mortality of early life stages and subsequent decreases in densities of adult females. Flow management that better mimics natural flow timing is likely to promote persistence of these species and others with similar phenology.     

Antagonistic effect of natural habitat conversion on community adjustment to climate warming in nonbreeding waterbirds

Although the impacts of climate and land-use changes on biodiversity have been widely documented, their joint effects remain poorly understood. We evaluated how nonbreeding waterbird communities adjust to climate warming along a gradient of land-use change. Using midwinter waterbird counts (132 species) at 164 major nonbreeding sites in 22 Mediterranean countries, we assessed the changes in species composition from 1991 to 2010, relative to thermal niche position and breadth, in response to regional and local winter temperature anomalies and conversion of natural habitats. We observed a low-level, nonsignificant community adjustment to the temperature increase where natural habitat conversion occurred. At the sites affected by natural habitat conversion, the relative increase of warm-dwelling species in response to climate warming was 6 times lower and the relative species decline was 3 times higher than in the sites without natural habitat conversion. We found no evidence of community adjustment to climate warming when natural habitat conversion was >5% over 15 years. This strong negative effect suggests an antagonistic interaction between climate warming and habitat change. These results underline the importance of habitat conservation in community adjustment to climate warming.     

Testis size variation in frogs: testing the alternatives

While sperm competition may be a major factor affecting relative testis size in vertebrates as a whole, additional hypotheses have not been given much attention in frogs. This is important because sperm competition is relatively uncommon in frogs and relative testis size varies in frogs that do not have multiple-male breeding systems. This paper tests two additional hypotheses for differences in relative testis size among frogs: relative clutch size (number of eggs/snout vent length) and androgen level. Testis size was measured in 90 species of frogs belonging to five families. Relative testis size was found to be positively correlated with relative clutch size in species that lack sperm competition. Mean androgen levels of species also positively covaried with relative testis size. However, there was no correlation between relative testis size and level of male agonistic behavior among species, despite other work indicating that testosterone levels are positively correlated with agonistic behavior in at least some species. These findings suggest that a number of factors in addition to sperm competition are important in the evolution of testis size in male frogs. Received: 17 January 1997 / Accepted after revision: 23 June 1997     

Habitat use and group size of pied cormorants (<Emphasis Type="Italic">Phalacrocorax varius</Emphasis>) in a seagrass ecosystem: possible effects of food abundance and predation risk

Both food abundance and predation risk may influence habitat use decisions. However, studies of habitat use by birds in marine environments have focused only on food abundance. I investigated the possible influences of food abundance and predation risk from tiger sharks (Galeocerdo cuvier) on habitat use by pied cormorants (Phalacrocorax varius) over two spatial scales and on cormorant group size. Cormorants were usually solitary, but group size was highest in shallow habitats during months when shark density was low. Regardless of season, cormorant density within shallow habitats was higher over seagrass than sand, and cormorants were distributed between these two microhabitats proportional to prey density. Therefore, cormorants appear to respond to prey abundance at a relatively narrow spatial scale (i.e., tens of meters). At the habitat-patch scale (~1 km), the density of cormorants and their prey (teleosts) was higher in shallow habitats than in deep ones, but the density of cormorants was influenced by an interaction between water temperature (i.e., season) and habitat. There was decreased use of shallow habitats as water temperature, and the density of tiger sharks, increased. When shark density was low, cormorants were distributed across habitats roughly in proportion to the abundance of fish, suggesting that cormorants respond to food abundance at the scale of habitat patches. However, as shark abundance increased, the relative density of cormorants dropped in the dangerous shallow habitats such that there was a greater density of cormorants relative to their food in deep habitats when sharks were abundant. This suggests that pied cormorants trade-off food and risk by accepting lower energetic returns to forage in safer habitats. This study provides the first evidence that marine habitat selection by birds may be influenced by such a trade-off, and provides further evidence that tiger sharks are important in determining habitat use of their prey and mediating indirect interactions within Shark Bay.Communicated by P. W. Sammarco, Chauvin     

Cryptic biodiversity effects: importance of functional redundancy revealed through addition of food web complexity

Interactions between predators and the degree of functional redundancy among multiple predator species may determine whether herbivores experience increased or decreased predation risk. Specialist parasites can modify predator behavior, yet rarely have cascading effects on multiple predator species and prey been evaluated. We examined influences of specialist phorid parasites (Pseudacteon spp.) on three predatory ant species and herbivores in a coffee agroecosystem. Specifically, we examined whether changes in ant richness affected fruit damage by the coffee berry borer (Hypothenemus hampei) and whether phorids altered multi-predator effects. Each ant species reduced borer damage, and without phorids, increasing predator richness did not further decrease borer damage. However, with phorids, activity of one ant species was reduced, indicating that the presence of multiple ant species was necessary to limit borer damage. In addition, phorid presence revealed synergistic effects of multiple ant species, not observed without the presence of this parasite. Thus, a trait-mediated cascade resulting from a parasite-induced predator behavioral change revealed the importance of functional redundancy, predator diversity, and food web complexity for control of this important pest.