Cover Caption

Cover : Diamondback terrapins (Malaclemys terrapin) inhabit brackish waters throughout the Atlantic and Gulf coasts of the United States. In parts of the turtle's range, juveniles (pictured) and adult males that inadvertently enter commercial crab traps typically drown, whereas the larger carapace of adult females excludes them from traps. On pages 1268‐1277, Wolak et al. document sex‐speci. c changes in carapace length and width in populations that co‐occur with commercial crab traps. Th ese changes have not been observed in contemporary and historic populations in areas without crab trapping.     

Diamondback Terrapin (Malaclemys terrapin) Mortality in Crab Pots

The entrapment of diamondback terrapins (  Malaclemys terrapin) in crab pots frequently results in drowning and death of the trapped turtles. We determined the rate of capture, size, sex, and age of terrapins captured, and the potential impact crab pot mortality has on local terrapin populations. We estimated terrapin capture rates of 0.17 terrapins/pot/day in shallow water areas of Chesapeake Bay (Maryland, USA). The sex ratio of terrapins caught in crab pots was 3:2 male biased because female terrapins become too large to enter crab pots by the time they reach 8 years of age. Males, however, remain vulnerable to entrapment throughout their life. Our estimates of capture rates and local population size suggest that 15–78% of a local population may be captured in a single year. As a consequence, crab pots may be the major reason terrapins are extirpated in coastal, shallow water areas with heavy crab pot fisheries. Additionally, the selective removal of males may also contribute to female-biased sex ratios observed in this diamondback terrapin population. We developed and tested a modified crab pot that increases terrapin survival and does not reduce the number of crabs caught. Our modified crab pot maintained permanent access to air and prevented the drowning of terrapins. Additionally, our modified crab pot caught more crabs than standard commercial crab pots, suggesting that the modified crab pot could be a viable alternative to standard traps that result in terrapin mortality.     

Relationships between sponges and crabs: patterns of epibiosis on Inachus aguiarii (Decapoda: Majidae)

The association between sponges and the crab Inachus aguiarii Brito Capello, 1876 was studied by analysing the relationships between sponge distributional patterns on the crab carapaces and several morphological and biological crab parameters. Juveniles, mature females and mature males were differentiated on the basis of sex dimorphism and terminal pubertary moult. All three groups were fouled to different degrees by sponges. Percent sponge cover was related to carapace size only in mature males whereas mature females, all but one of which were ovigerous, were extensively covered regard-less of their size. It is proposed that some behavioural patterns unique to females, such as long resting periods in sponge-rich microhabitats, are responsible for these high sponge covers in females. Sponges showed two trends in the colonization of the carapace, leading to either a monopolizing or a sharing of the available carapace surface. The sponge species found on the carapaces studied are not obligatory epibionts of crabs, but are believed to reflect the sponge population characterizing the crab home range.     

Using Parasitic Trematode Larvae to Quantify an Elusive Vertebrate Host

Abstract: Digenean trematode parasites require multiple host species to complete their life cycles, and their abundance can often be strongly correlated with the abundance of their host species. Species richness and abundance of parasites in easily sampled host species may yield an accurate estimate of the species richness and abundance of other hosts in a parasite's life cycle that are difficult to survey directly. Accordingly, we investigated whether prevalence and mean abundance of trematodes could be used to estimate the abundance of one of their host species, diamondback terrapins (Malaclemys terrapin), which are difficult to sample and are designated as near threatened (by the International Union for Conservation of Nature [IUCN Red List]) along some U.S. coasts. As an adult the trematode Pleurogonius malaclemys is specific to terrapins. Its larval stages live first inside mud snails (Ilyanassa obsoleta) and are subsequently shed into the environment where they form external metacercarial cysts on hard surfaces such as snail opercula. The life cycle of P. malaclemys is completed when terrapins ingest these cysts. At 12 sites along the coast of Georgia (U.S.A.), we determined the prevalence of internal P. malaclemys larvae in mud snails (proportion of infected snails in a population) and the prevalence and mean abundance of external trematode cysts. We examined whether these data were correlated with terrapin abundance, which we estimated with mark‐recapture methods. The abundance of external cysts and salinity explained ≥59% of the variability in terrapin abundance. We suggest that dependent linkages between the life stages of multihost parasites make them reliable predictors of host species’ abundance, including hosts with abundances that are challenging to quantify directly.     

Use of Fully Biodegradable Panels to Reduce Derelict Pot Threats to Marine Fauna

Fishing pots (i.e., traps) are designed to catch fish or crustaceans and are used globally. Lost pots are a concern for a variety of fisheries, and there are reports that 10 – 70% of deployed pots are lost annually. Derelict fishing pots can be a source of mortality for target and bycatch species for several years. Because continual removal of derelict gear can be impractical over large spatial extents, modifications are needed to disarm gear once it is lost. We tested a fully biodegradable panel with a cull or escape ring designed for placement on the sides of a crab pot that completely degrades into environmentally neutral constituents after approximately 1 year. This panel is relatively inexpensive, easy to install, and can be used in multiple fisheries. We used the blue crab (Callinectes sapidus) fishery as a test case because it is a large pot fishery and blue crab pots are similar to traps used in other pot fisheries. We had commercial fishers deploy pots with panels alongside standard pots in Chesapeake Bay (U.S.A.) to assess potential effects of our experimental pots on blue crab catch. We compared the number, biomass, and size of crabs captured between standard and experimental pots and evaluated differences in catch over a crabbing season (March–November) at five locations. There was no evidence that biodegradable panels adversely affected catch. In all locations and time periods, legal catches were comparable in abundance, biomass, and size between experimental and standard pots. Properly designed biodegradable panels appear to be a viable solution to mitigate adverse effects of derelict pots. Uso de Paneles Completamente Biodegradables para Reducir las Amenazas de Vasijas Abandonadas para la Fauna Marina     

Male Mutation Bias and Possible Long‐Term Effects of Human Activities

Abstract: The ability of a population to adapt to changing environments depends critically on the amount and kind of genetic variability it possesses. Mutations are an important source of new genetic variability and may lead to new adaptations, especially if the population size is large. Mutation rates are extremely variable between and within species, and males usually have higher mutation rates as a result of elevated rates of male germ cell division. This male bias affects the overall mutation rate. We examined the factors that influence male mutation bias, and focused on the effects of classical life‐history parameters, such as the average age at reproduction and elevated rates of sperm production in response to sexual selection and sperm competition. We argue that human‐induced changes in age at reproduction or in sexual selection will affect male mutation biases and hence overall mutation rates. Depending on the effective population size, these changes are likely to influence the long‐term persistence of a population.     

Breeding Periodicity for Male Sea Turtles,Operational Sex Ratios,and Implications in the Face of Climate Change

Abstract: Species that have temperature‐dependent sex determination (TSD) often produce highly skewed offspring sex ratios contrary to long‐standing theoretical predictions. This ecological enigma has provoked concern that climate change may induce the production of single‐sex generations and hence lead to population extirpation. All species of sea turtles exhibit TSD, many are already endangered, and most already produce sex ratios skewed to the sex produced at warmer temperatures (females). We tracked male loggerhead turtles (Caretta caretta) from Zakynthos, Greece, throughout the entire interval between successive breeding seasons and identified individuals on their breeding grounds, using photoidentification, to determine breeding periodicity and operational sex ratios. Males returned to breed at least twice as frequently as females. We estimated that the hatchling sex ratio of 70:30 female to male for this rookery will translate into an overall operational sex ratio (OSR) (i.e., ratio of total number of males vs females breeding each year) of close to 50:50 female to male. We followed three male turtles for between 10 and 12 months during which time they all traveled back to the breeding grounds. Flipper tagging revealed the proportion of females returning to nest after intervals of 1, 2, 3, and 4 years were 0.21, 0.38, 0.29, and 0.12, respectively (mean interval 2.3 years). A further nine male turtles were tracked for short periods to determine their departure date from the breeding grounds. These departure dates were combined with a photoidentification data set of 165 individuals identified on in‐water transect surveys at the start of the breeding season to develop a statistical model of the population dynamics. This model produced a maximum likelihood estimate that males visit the breeding site 2.6 times more often than females (95%CI 2.1, 3.1), which was consistent with the data from satellite tracking and flipper tagging. Increased frequency of male breeding will help ameliorate female‐biased hatchling sex ratios. Combined with the ability of males to fertilize the eggs of many females and for females to store sperm to fertilize many clutches, our results imply that effects of climate change on the viability of sea turtle populations are likely to be less acute than previously suspected.     

Reexamining the Minimum Viable Population Concept for Long‐Lived Species

For decades conservation biologists have proposed general rules of thumb for minimum viable population size (MVP); typically, they range from hundreds to thousands of individuals. These rules have shifted conservation resources away from small and fragmented populations. We examined whether iteroparous, long‐lived species might constitute an exception to general MVP guidelines. On the basis of results from a 10‐year capture‐recapture study in eastern New York (U.S.A.), we developed a comprehensive demographic model for the globally threatened bog turtle (Glyptemys muhlenbergii), which is designated as endangered by the IUCN in 2011. We assessed population viability across a wide range of initial abundances and carrying capacities. Not accounting for inbreeding, our results suggest that bog turtle colonies with as few as 15 breeding females have >90% probability of persisting for >100 years, provided vital rates and environmental variance remain at currently estimated levels. On the basis of our results, we suggest that MVP thresholds may be 1–2 orders of magnitude too high for many long‐lived organisms. Consequently, protection of small and fragmented populations may constitute a viable conservation option for such species, especially in a regional or metapopulation context. Reexaminando el Concepto de Población Mínima Viable para Especies Longevas Resumen     

Sexual dimorphism in the sand dollar Echinarachnius parma

Samples of Echinarachnius parma (Lamarck), collected during summer 1988 from three sites along the eastern coast of Canada (the northern Gulf of St. Lawrence, Chaleur Bay and the Bay of Fundy), showed sexual dimorphism in the shape of the oral surface in addition to marked changes in the camber of the aboral surface related to the strength of wave and current activity. The oral surface is more concave in males than in females. The dimorphism develops as individuals become sexually mature and is pronounced for individuals measuring >4 cm in diameter. Associated with this shape difference, the volume of females is greater than for males, and this corresponds to the larger size attained by the ovaries compared to the testis. Shape differences between males and females are rare in echinoderms, and E. parma is the first species for which the sexes can be separated with a high degree of accuracy (>80%) on the basis of shape.     

Juvenile length-at-age data reveal that spanner crabs (<Emphasis Type="Italic">Ranina ranina</Emphasis>) grow slowly

The spanner crab (Ranina ranina) is a widespread and abundant brachyuran in offshore sand substrata of the Indo-Pacific region. Little is known of this species biology, population dynamics and ecology, despite it being the target of commercial fishing operations in many areas. Previous studies of R. ranina growth using length-frequency analysis of samples collected with commercial fishing gear have derived widely divergent estimates of growth parameters. The estimated time taken to reach 100 mm rostral carapace length (minimum legally exploited size in Queensland, Australia) in those studies has ranged from 1.75 to 8.83 years for females and from 1.08 to 3.58 years for males. Our data show that the commercial fishing apparatus used in those studies is size selective and catches only adult crabs. The resulting size bias in samples collected using that apparatus precludes the application of length-frequency-based techniques to estimate growth parameters from those samples. We devised a new dredge to collect samples of juvenile R. ranina and to calculate juvenile growth rates from modal progression in those samples. We combined those data with estimated mean maximum lengths (L) of 121.7 mm for females and 155.9 mm for males from commercial catch data to model other von Bertalanffy growth parameters using bootstrap methods. Those modelled parameters (K=0.29, T₀=–0.24 for females; K=0.23, T₀=–0.25 for males) indicate that R. ranina grows more slowly than most previous estimates suggest, with females requiring an average of 6.35 years and males 4.31 years to reach 100 mm rostral carapace length. This slow growth is consistent with the slow metabolism of R. ranina, and indicates that this species would be likely to recover slowly from overexploitation.Communicated by G.F. Humphrey, Sydney     

Pheromone Monitoring of Rare and Threatened Insects: Exploiting a Pheromone–Kairomone System to Estimate Prey and Predator Abundance

Abstract: Pheromone‐based monitoring is a promising new method for assessing the conservation status of many threatened insect species. We examined the versatility and usefulness of pheromone‐based monitoring by integrating a pheromone–kairomone trapping system and pitfall trapping system in the monitoring of two saproxylic beetles, the hermit beetle Osmoderma eremita (Coleoptera: Scarabaeidae) and its predator Elater ferrugineus (Coleoptera: Elateridae), which live inside hollow trees. We performed mark–recapture studies of both species with unbaited pitfall traps in oak hollows combined with pheromone‐baited funnel traps suspended from oak branches to intercept dispersing individuals. For O. eremita, the integrated trapping system showed that the population in the study sites may be considerably higher than estimates based on extrapolation from pitfall trapping alone (approximately 3400 vs. 1100 or 1800 individuals, respectively). Recaptures between odor‐baited funnel traps showed that males and females had similar dispersal rates, but estimating the number of dispersing individuals was problematic due to declining recapture probability between subsequent capture events. Our conservative estimate, assuming a linear decrease in capture probability, suggested that around 1900 individuals, or at least half of the O. eremita population, may perform flights from their natal host trees, representing higher dispersal rates than previous estimates. E. ferrugineus was rarely caught in pitfall traps. One hundred thirty‐nine individuals, likely almost exclusively females, were caught in odor‐baited funnel traps with approximately 4% recapture probability. If recapture probability over consecutive capture events follows that of O. eremita, this would correspond to a total population size of 2500–3000 individuals of the predator; similar to its supposed prey O. eremita. Our results demonstrate that pheromone‐based monitoring is a valuable tool in the study of species or life‐history stages that would otherwise be inaccessible.     

Effectiveness of Predator Removal for Enhancing Bird Populations

Abstract: Predation pressure on vulnerable bird species has made predator control an important issue for international nature conservation. Predator removal by culling or translocation is controversial, expensive, and time‐consuming, and results are often temporary. Thus, it is important to assess its effectiveness from all available evidence. We used explicit systematic review methodology to determine the impact of predator removal on four measurable responses in birds: breeding performance (hatching success and fledging success) and population size (breeding and postbreeding). We used meta‐analysis to summarize results from 83 predator removal studies from six continents. We also investigated whether characteristics of the prey, predator species, location, and study methodology explained heterogeneity in effect sizes. Removing predators increased hatching success, fledging success, and breeding populations. Removing all predator species achieved a significantly larger increase in breeding population than removing only a subset. Postbreeding population size was not improved on islands, or overall, but did increase on mainlands. Heterogeneity in effect sizes for the four population parameters was not explained by whether predators were native or introduced; prey were declining, migratory, or game species; or by the study methodology. Effect sizes for fledging success were smaller for ground‐nesting birds than those that nest elsewhere, but the difference was not significant. We conclude that current evidence indicates that predator removal is an effective strategy for the conservation of vulnerable bird populations. Nevertheless, the ethical and practical problems associated with predator removal may lead managers to favor alternative, nonlethal solutions. Research is needed to provide and synthesize data to determine whether these are effective management practices for future policies on bird conservation.     

Conservation genomics of the endangered Burmese roofed turtle

The Burmese roofed turtle (Batagur trivittata) is one of the world's most endangered turtles. Only one wild population remains in Myanmar. There are thought to be 12 breeding turtles in the wild. Conservation efforts for the species have raised >700 captive turtles since 2002, predominantly from eggs collected in the wild. We collected tissue samples from 445 individuals (approximately 40% of the turtles’ remaining global population), applied double‐digest restriction‐site associated DNA sequencing (ddRAD‐Seq), and obtained approximately 1500 unlinked genome‐wide single nucleotide polymorphisms. Individuals fell into 5 distinct genetic clusters, 4 of which represented full‐sib families. We inferred a low effective population size (≤10 individuals) but did not detect signs of severe inbreeding, possibly because the population bottleneck occurred recently. Two groups of 30 individuals from the captive pool that were the most genetically diverse were reintroduced to the wild, leading to an increase in the number of fertile eggs (n = 27) in the wild. Another 25 individuals, selected based on the same criteria, were transferred to the Singapore Zoo as an assurance colony. Our study demonstrates that the research‐to‐application gap in conservation can be bridged through application of cutting‐edge genomic methods.     

Reducing the Maladaptive Attractiveness of Solar Panels to Polarotactic Insects

Abstract: Human‐made objects (e.g., buildings with glass surfaces) can reflect horizontally polarized light so strongly that they appear to aquatic insects to be bodies of water. Insects that lay eggs in water are especially attracted to such structures because these insects use horizontal polarization of light off bodies of water to find egg‐laying sites. Thus, these sources of polarized light can become ecological traps associated with reproductive failure and mortality in organisms that are attracted to them and by extension with rapid population declines or collapse. Solar panels are a new source of polarized light pollution. Using imaging polarimetry, we measured the reflection–polarization characteristics of different solar panels and in multiple‐choice experiments in the field we tested their attractiveness to mayflies, caddis flies, dolichopodids, and tabanids. At the Brewster angle, solar panels polarized reflected light almost completely (degree of polarization d ≈100%) and substantially exceeded typical polarization values for water (d ≈30–70%). Mayflies (Ephemeroptera), stoneflies (Trichoptera), dolichopodid dipterans, and tabanid flies (Tabanidae) were the most attracted to solar panels and exhibited oviposition behavior above solar panels more often than above surfaces with lower degrees of polarization (including water), but in general they avoided solar cells with nonpolarizing white borders and white grates. The highly and horizontally polarizing surfaces that had nonpolarizing, white cell borders were 10‐ to 26‐fold less attractive to insects than the same panels without white partitions. Although solar panels can act as ecological traps, fragmenting their solar‐active area does lessen their attractiveness to polarotactic insects. The design of solar panels and collectors and their placement relative to aquatic habitats will likely affect populations of aquatic insects that use polarized light as a behavioral cue.     

Role of Familiarity and Preference in Reproductive Success in Ex Situ Breeding Programs

Abstract: Success of captive‐breeding programs centers on consistent reproduction among captive animals. However, many individuals do not reproduce even when they are apparently healthy and presented with mates. Mate choice can affect multiple parameters of reproductive success, including mating success, offspring production, offspring survival, and offspring fecundity. We investigated the role of familiarity and preference on reproductive success of female Columbia Basin pygmy rabbits (Brachylagus idahoensis) as measured by litter production, litter size, average number of young that emerged from the burrow, and average number of young that survived to 1 year. We conducted these studies on pygmy rabbits at the Oregon Zoo (Portland, Oregon, U.S.A.) and Washington State University (Pullman, Washington, U.S.A.) from February to June 2006, 2007, and 2008. Before mating, we housed each female adjacent to 2 males (neighbors). Female preference for each potential mate was determined on the basis of behavioral interactions observed and measured between the rabbits. We compared reproductive success between females mated with neighbor and non‐neighbor males and between females mated with preferred and nonpreferred males. Our findings suggest that mating with a neighbor compared with a non‐neighbor and mating with a preferred neighbor compared with a nonpreferred neighbor increased reproductive success in female pygmy rabbits. Litter production, average number of young that emerged, and average number of young that survived to 1 year were higher in rabbits that were neighbors before mating than in animals who were not neighbors. Pairing rabbits with a preferred partner increased the probability of producing a litter and was significantly associated with increased litter size. In captive breeding programs, mates are traditionally selected on the basis of genetic parameters to minimize loss of genetic diversity and inbreeding coefficients. Our results suggest that integrating genetic information with social dynamics and behavioral measures of preference may increase the reproductive output of the pygmy rabbit captive‐breeding program. Our findings are consistent with the idea that allowing mate choice and familiarity increase the reproductive success of captive‐breeding programs for endangered species.     

Effects of Forest Fragmentation on Seed Dispersal and Seedling Establishment in Ornithochorous Trees

Abstract: Habitat fragmentation increases seed dispersal limitation across the landscape and may also affect subsequent demographic stages such as seedling establishment. Thus, the development of adequate plans for forest restoration requires an understanding of mechanisms by which fragmentation hampers seed delivery to deforested areas and knowledge of how fragmentation affects the relationship between seed‐deposition patterns and seedling establishment. We evaluated the dispersal and recruitment of two bird‐dispersed, fleshy‐fruited tree species (Crataegus monogyna and Ilex aquifolium) in fragmented secondary forests of northern Spain. Forest fragmentation reduced the probability of seed deposition for both trees because of decreased availability of woody perches and fruit‐rich neighborhoods for seed dispersers, rather than because of reductions in tree cover by itself. The effects of fragmentation went beyond effects on the dispersal stage in Crataegus because seedling establishment was proportional to the quantities of bird‐dispersed seeds arriving at microsites. In contrast, postdispersal mortality in Ilex was so high that it obscured the seed‐to‐seedling transition. These results suggest that the effects of fragmentation are not necessarily consistent across stages of recruitment across species. Habitat management seeking to overcome barriers to forest recovery must include the preservation, and even the planting, of fleshy‐fruited trees in the unforested matrix as a measure to encourage frugivorous birds to enter into open and degraded areas. An integrative management strategy should also explicitly consider seed‐survival expectancies at microhabitats to preserve plant‐population dynamics and community structure in fragmented landscapes.     

Estimation of Body‐Size Traits by Photogrammetry in Large Mammals to Inform Conservation

Abstract: Photography, including remote imagery and camera traps, has contributed substantially to conservation. However, the potential to use photography to understand demography and inform policy is limited. To have practical value, remote assessments must be reasonably accurate and widely deployable. Prior efforts to develop noninvasive methods of estimating trait size have been motivated by a desire to answer evolutionary questions, measure physiological growth, or, in the case of illegal trade, assess economics of horn sizes; but rarely have such methods been directed at conservation. Here I demonstrate a simple, noninvasive photographic technique and address how knowledge of values of individual‐specific metrics bears on conservation policy. I used 10 years of data on juvenile moose (Alces alces) to examine whether body size and probability of survival are positively correlated in cold climates. I investigated whether the presence of mothers improved juvenile survival. The posited latter relation is relevant to policy because harvest of adult females has been permitted in some Canadian and American jurisdictions under the assumption that probability of survival of young is independent of maternal presence. The accuracy of estimates of head sizes made from photographs exceeded 98%. The estimates revealed that overwinter juvenile survival had no relation to the juvenile's estimated mass (p < 0.64) and was more strongly associated with maternal presence (p < 0.02) than winter snow depth (p < 0.18). These findings highlight the effects on survival of a social dynamic (the mother‐young association) rather than body size and suggest a change in harvest policy will increase survival. Furthermore, photographic imaging of growth of individual juvenile muskoxen (Ovibos moschatus) over 3 Arctic winters revealed annual variability in size, which supports the idea that noninvasive monitoring may allow one to detect how some environmental conditions ultimately affect body growth.     

Effects of Multiple Levels of Social Organization on Survival and Abundance

Abstract: Identifying how social organization shapes individual behavior, survival, and fecundity of animals that live in groups can inform conservation efforts and improve forecasts of population abundance, even when the mechanism responsible for group‐level differences is unknown. We constructed a hierarchical Bayesian model to quantify the relative variability in survival rates among different levels of social organization (matrilines and pods) of an endangered population of killer whales (Orcinus orca). Individual killer whales often participate in group activities such as prey sharing and cooperative hunting. The estimated age‐specific survival probabilities and survivorship curves differed considerably among pods and to a lesser extent among matrilines (within pods). Across all pods, males had lower life expectancy than females. Differences in survival between pods may be caused by a combination of factors that vary across the population's range, including reduced prey availability, contaminants in prey, and human activity. Our modeling approach could be applied to demographic rates for other species and for parameters other than survival, including reproduction, prey selection, movement, and detection probabilities.     

Ecological Trap for Desert Lizards Caused by Anthropogenic Changes in Habitat Structure that Favor Predator Activity

Abstract: Anthropogenic habitat perturbation is a major cause of population decline. A standard practice managers use to protect populations is to leave portions of natural habitat intact. We describe a case study in which, despite the use of this practice, the critically endangered lizard Acanthodactylus beershebensis was locally extirpated from both manipulated and natural patches within a mosaic landscape of an afforestation project. We hypothesized that increased structural complexity in planted patches favors avian predator activity and makes these patches less suitable for lizards due to a heightened risk of predation. Spatial rarity of natural perches (e.g., trees) in arid scrublands may hinder the ability of desert lizards to associate perches with low‐quality habitat, turning planted patches into ecological traps for such species. We erected artificial trees in a structurally simple arid habitat (similar to the way trees were planted in the afforestation project) and compared lizard population dynamics in plots with these structures and without. Survival of lizards in the plots with artificial trees was lower than survival in plots without artificial trees. Hatchlings dispersed into plots with artificial trees in a manner that indicated they perceived the quality of these plots as similar to the surrounding, unmanipulated landscape. Our results showed that local anthropogenic changes in habitat structure that seem relatively harmless may have a considerable negative effect beyond the immediate area of the perturbation because the disturbed habitat may become an ecological trap.