Experimental analysis of diet specialization in the snail kite: the role of behavioral conservatism

We examined factors maintaining extreme diet specialization in the snail kite (Rostrhamus sociabilis), a medium-sized hawk which feed almost exclusively on Pomacea snails, by determining why during some months kites eat crabs (Dilocarcinus dentatus) in the Ilanos of Venezuela. We offered snails and crabs of different sizes to wild free-flying birds to develop estimates for a prey choice model. Handling times of Pomacea doliodes snails averaged 90±39 s and were positively correlated with snail size. Handling times for crabs (x=353±130 s) were significantly longer and exhibited greater variation than for snails, and were not correlated with crab size. Edible crab tissues had greater dry weights and contained more energy (25.37 kJ/g) than tissues of snails (16.91 kJ/g). Total energy of crabs was much greater than that of snails, and total energy of both foods was highly related to body length. We constructed an allometric equation for profitability of snails and crabs. Snails were more profitable than all but the largest crabs, but estimates of variance in profitability were greater for crabs. Predictions from the model were tested by offering crabs that represented equal, greater and much greater profitability than snails, to determine whether kites chose prey according to profitability. Only 15.6% of 289 food items chosen were crabs. Half of the 18 kites tested did not eat crabs and only 3 birds switched from snails to more profitable crabs. Four fledglings showed no preference for snails. The role of neophobia in food choice was investigated by offering unfamiliar snails (Pomacea urceus) to kites. Kites exhibited neophobic behaviors, and 5 of 12 birds chose not to capture P. urceus. Two-thirds of the 12 snails chosen were rejected immediately, but the others were handled efficiently (x=133±89 s). Although morphological adaptations allow kites to specialize on snails, the costs of specialization were overcome for kites when the profitability of alternative food increased sufficiently. Our results suggest a role for behavioral conservatism, in the form of risk-averse foraging and neophobia, in maintaining severe diet specialization in the snail kite.     

Experimental Test of Preferences for an Invasive Prey by an Endangered Predator: Implications for Conservation

Identifying impacts of exotic species on native populations is central to ecology and conservation. Although the effects of exotic predators on native prey have received much attention, the role of exotic prey on native predators is poorly understood. Determining if native predators actively prefer invasive prey over native prey has implications for interpreting invasion impacts, identifying the presence of evolutionary traps, and predator persistence. One of the world’s most invasive species, Pomacea maculata, has recently established in portions of the endangered Everglade snail kite’s (Rostrhamus sociabilis plumbeus) geographic range. Although these exotic snails could provide additional prey resources, they are typically much larger than the native snail, which can lead to lower foraging success and the potential for diminished energetic benefits in comparison to native snails. Nonetheless, snail kites frequently forage on exotic snails. We used choice experiments to evaluate snail kite foraging preference in relation to exotic species and snail size. We found that snail kites do not show a preference for native or exotic snails. Rather, snail kites generally showed a preference for medium-sized snails, the sizes reflective of large native snails. These results suggest that while snail kites frequently forage on exotic snails in the wild, this behavior is likely driven simply by the abundance of exotic snails rather than snail kites preferring exotics. This lack of preference offers insights to hypotheses regarding effects of exotic species, guidance regarding habitat and invasive species management, and illustrates how native-exotic relationships can be misleading in the absence of experimental tests of such interactions.     

Natural selection by avian predators on size and colour of a freshwater snail (Pomacea flagellatd)

We identify two avian predators of the Neotropical apple snail, Pomaceaflagellata, and estimate the strength, direction and form of multivariate natural selection by these predators on size and colour of snail shells. Limpkins are tactile predators and act as agents of disruptive selection on snail size, selecting average-sized snails disproportionately more often than small or large snails (y = 0.39, SE = 0.08). In addition, we were able to identify variation in handling behaviours and snail size selection among individual limpkins. Individual limpkins showed preferences for snails of different sizes and punctured the snail shells opposite the aperture mainly when handling large snails. Snail kites are visual predators and seem to be agents of directional selection against lighter coloured snails (β= 0.66, SE = 0.33). The ecological interaction between the apple snail and its predators provides a powerful system to further explore the role of predation in determining evolutionary changes in snail behaviour, morphology and life history.     

Multiscale patterns of movement in fragmented landscapes and consequences on demography of the snail kite in Florida

1. Habitat loss and fragmentation are major factors affecting vertebrate populations. A major effect of these habitat alterations is that they reduce movement of organisms. Despite the accepted importance of movement in driving the dynamics of many natural populations, movement of vertebrates in fragmented landscapes have seldom been estimated with robust statistical methods. 2. We estimated movement probabilities of snail kites Rosthramus sociabilis within the remaining wetlands in Florida. Using both radio-telemetry and banding information, we used a multistate modelling approach to estimate transition probabilities at two temporal scales (month; year) and multiple spatial scales. We examined kite movement among wetlands altered by three different levels of fragmentation: among wetlands separated by small physical barriers (e.g. road); among wetlands separated by moderate amount of matrix (< 5 km); and among wetlands separated by extensive matrix areas (> 15 km). 3. Kites moved extensively among contiguous wetlands (movement probability 0.29 per month), but significantly less among isolated wetlands (movement probability 0.10 per month). 4. Kites showed high levels of annual site fidelity to most isolated wetlands (probability ranged from 0.72 to 0.95 per year). 5. We tested the effects of patch size and interpatch distance on movement. Our modelling indicated an effect of both distance and patch size on juveniles' movement (but not adult) when examining movements among fragments. 6. Only a small proportion of kites escaped a regional drought by moving to refugia (wetlands less affected by drought). Many individuals died after the drought. During drought adult survival dropped by 16% while juvenile survival dropped by 86% (possibly because juveniles were less likely to reach refugia). 7. We hypothesize that fragmentation may decrease kite's resistance to drought by restricting exploratory behaviour.     

Crayfish predation on the common pond snail (Lymnaea stagnalis): the effect of habitat complexity and snail size on foraging efficiency

Optimal foraging theory was used to explain selective foraging by the introduced signal crayfish (Pacifastacus leniusculus) on the thin-shelled common pond snail (Lymnaea stagnalis). Crayfish predation efficiency was studied in relation to habitat complexity and snail size. In a pool experiment (area 1.3 m²) single adult crayfish were allowed to feed on four size classes of snails for one week. A pair-wise preference trial (aquarium experiment) tested if adult crayfish selectively predated on particular size classes of snail and if prey value (expressed as snail dry mass per handling time) could explain the size range of snails chosen. Crayfish preferred the smallest size classes of snails in both pool and aquaria experiments. In the pool experiment crayfish had a strong effect on snail survival. Habitat complexity did not affect overall snail survival, but resulted in reduced predation pressure on the smallest size classes of snails. Handling time and shell-thickness increased exponentially with increasing snail size, and the two smallest size classes had the highest prey values. The results suggest that crayfish can structure the abundance and size distribution of thin-shelled snails, through size-selective predation and reduction of macrophytes. The mechanisms behind the choice of snails may be based on prey value and reduced exposure time to predators and conspecifics. Crayfish effects on snail size distribution may be less pronounced in complex habitats such as macrophyte beds. This revised version was published online in July 2006 with corrections to the Cover Date.     

Habitat selection of the Black-shouldered Kite Elanus caeruleus in agroecosystems of Swaziland

A decline in many African raptor populations may contribute to loss of ecosystem services such as pest control. The Black-shouldered Kite Elanus caeruleus stands out as an exception, experiencing range expansions due to its proclivity for agricultural landscapes. We surveyed roads throughout agroecosystems in Swaziland for Black-shouldered Kites and looked at habitat selection in a used versus available framework. We found that kites selected for savannas and tall herbaceous vegetation but did not show preferences for or against land-use types. We also found a significant relationship with kite presence and low tree densities in the immediate vicinity of a perch site. Singles, pairs and fledgling kites were observed in agricultural landscapes, suggesting that although they are not selecting for these areas in proportion to their availability, they are foraging in it. This may be a boon for farmers who implement ecologically based rodent management programs.     

Prioritizing species conservation: does the Cape Verde kite exist?

The Cape Verde kite (Milvus milvus fasciicauda) is considered to be one of the rarest birds of prey in the world and at significant risk of extinction. For this reason there is great interest in both the taxonomic and the population status of this group. To help resolve its taxonomic status, we provide phylogenetic analyses based on three mitochondrial genes for a sampling of kites in the genus Milvus, including a broad geographical sampling of black kites (Milvus migrans), red kites (Milvus milvus), Cape Verde kite museum specimens collected between 1897 and 1924, and five kites trapped on the Cape Verde Islands during August 2002. We found that the historical Cape Verde kites, including the type specimen, were non-monophyletic and scattered within a larger red kite clade. The recently trapped kites from the Cape Verde Islands were all phylogenetically diagnosed as black kites. Our findings suggest that the traditional Cape Verde kite is not a distinctive evolutionary unit, and the case for species status, as recently suggested by others, is not supported. We do find support for recognition of at least one clade of yellow-billed kites, traditionally considered as a black kite subspecies, as a distinctive phylogenetic species.     

Space and habitat use by Red Kites Milvus milvus during winter in the Guadalquivir marshes: a comparison between resident and wintering populations

Differences in habitat selection, diet and behaviour of resident and wintering Red Kites Milvus milvus were studied in Donana National Park, southwest Spain. Adult resident Red Kites roosted at their nests, while immature residents and wintering birds gathered at communal roosts. Individuals remained on average for two consecutive nights (range 1–8) at the same communal roost. Wintering kites spent significantly more time foraging gregariously than residents. The use of the marsh was greater by wintering kites while residents more often used the forest. Wintering birds consumed more goose carrion than the residents. Wintering kites had larger core areas, moved farther from their roost sites to feed and changed foraging areas more frequently than residents. Among residents, adult females had the smallest home ranges (core area, distances travelled and time spent flying). We suggest that Red Kites wintering in Donana occupy the marsh because it provides high food availability, and is vacated by Black Kites M. migrans which exploit this profitable habitat during the breeding season.     

Communal Roosting of Wintering Red Kites Milvus milvus (Aves,Accipitridae): Social Feeding Strategies for the Exploitation of Food Resources

Social feeding strategies of wintering red kites are analyzed in relation to age, food, roost-sites and differences from kite residents. Whereas young and adult wintering kites gathered at roost sites almost daily, adult residents did not, and immature residents only occasionally. Kites using roost sites feed more often on prey prelocated by others, while lone roosters also forage and discover food alone. After finding food, kites tend to shift to a new roost site and foraging area. Two details of the ‘information centre’ hypothesis are confirmed in our study: carcasses are unpredictably found patches, divisible between several individuals. But carcasses disappeared fast in the study area, and no increase with time in the number of birds consuming a carcass was observed, so that information transmission was unconfirmed. When kites leave the roost in groups no leader is detectable. It seems that other types of social foraging are operating, and the model best matching our results is network foraging.     

Effects of body size and resource availability on dispersal in a native and a non-native estuarine snail

I manipulated snail densities of two coexisting species of salt marsh snail, Cerithidea californica Haldeman (native) and Batillaria attramentaria Sowerby (non-indigenous) to investigate how resource levels set by intraspecific competition may influence dispersal rates. I used two distinct size classes of the snails (mature and immature) to determine if the effects of competition on dispersal differed between developmental stages of a consumer. Dispersal attempts were measured within enclosure pens by counting snails climbing the sides of the enclosure. The influence of snail density per se and resource levels (which were set by snail densities) on dispersal rates were separated by comparing responses of snails to density before and after resources became depleted. For large snails of both species, dispersal increased as resource levels decreased, supporting the hypothesis that competition influences dispersal rates. Small snails of both species, in contrast, always dispersed at relatively higher rates than larger individuals, but were not influenced by variation in resource levels. This result corroborates other studies that have shown reduced competition in these species at smaller size, and suggests that another mechanism, such as genetically coded behavior to disperse when young, influences their behavior. Previous experiments demonstrated Batillaria's superior resource conversion efficiency; therefore, I had hypothesized that for any given resource level, Cerithidea would disperse more, because it was more affected by resource availability. Adult Batillaria, however, responded more sensitively to resource levels (i.e., dispersed more at any given resource level) than Cerithidea. This counter-intuitive result illustrates the potential importance of genetic limitations on behavioral responses available to a species. Constraints on behavioral responses may have been accentuated since Batillaria is a non-indigenous species whose evolved behavioral responses are not necessarily adapted to its present, non-native environment.     

Food partitioning between breeding White-tailed Kites (Elanus leucurus; Aves; Accipitridae) and Barn Owls (Tyto alba; Aves; Tytonidae) in southern Brazil.

I examined the diet of breeding White-tailed Kites (Elanus leucurus; Aves; Accipitridae) and Barn Owls (Tyto alba; Aves; Tytonidae) in an agrarian area of southern Brazil by analyzing regurgitated prey remains. The objective was to evaluate how these raptors, which differ markedly in their hunting activity periods (owls are nocturnal and kites diurnal), share their mammalian food component. 2,087 prey consumed by Barn Owls and 1,276 by White-tailed Kites were identified. They presented a high overlap of food-niches (Piankas index was 0.98). Based on the daily activity period of their main small mammal prey, a lower overlap would be expected. The crepuscular/nocturnal Mus musculus was the main prey for the diet of breeding Barn Owls (81%) and White-tailed Kites (63%). This small exotic rodent provided 63% of the small mammal biomass ingested by owls and 44% by kites. Larger native small mammals were also considered important for the diet of kites, mainly because of their biomass contribution. Although these raptors differ markedly in their hunting activity periods, Barn Owls and White-tailed Kites are very similar predators in southern Brazil, overlapping their diets.     

Physical and biological processes influencing zonation patterns of a subtidal population of the marine snail, Astraea (Lithopoma) undosa Wood 1828

The population density and size distribution of the marine gastropod, Astraea (Lithopoma) undosa Wood 1828, at Bird Rock, Santa Catalina Island, CA, reveal an inverse relationship between population density and mean individual size, over a depth gradient. This trend may be correlated with physical and biological differences between habitats for parameters such as water motion, competitive interactions, and predation.

The potential effect of hydrodynamic forces on the zonation patterns of Astraea undosa was tested in laboratory and field experiments. Based on theoretical predictions of the relationship between shear force and water velocity on different-sized snails, large snails are subjected to greater shear forces, as a result of water motion, than medium or small snails. Results of dislodgment experiments conducted in the laboratory indicated that for a given force per unit area, all snails dislodged at nearly the same frequency, with 50% of snails predicted to dislodge at about 4 m/s, and 100% of snails predicted to dislodge at about 8 m/s velocity. These results suggest that hydrodynamic forces may be an important factor in the shallowest subtidal zones.

A factorial-designed caging experiment was used to test the effects of snail population density on growth rates of snails of three different size classes. For small and medium size classes, results indicated an inverse relationship between population density and growth rates, which was especially pronounced for smaller snails. These data, in conjunction with long-term patterns of population density and size distribution in the field, suggest that intraspecific competition also plays a role in determining size-specific zonation patterns.

Tethering experiments, used to estimate predation rates in different algal-cover zones, suggest that there are no differences in survival rates among different snail size classes; however, survival rates differ among zones and may contribute further to the observed zonation patterns. Overall, data indicate that a combination of physical and biological processes controls the population density and size-distribution of Astraea undosa over a depth gradient at Santa Catalina Island, California.     

Affinity for natal environments by dispersers impacts reproduction and explains geographical structure of a highly mobile bird

Understanding dispersal and habitat selection behaviours is central to many problems in ecology, evolution and conservation. One factor often hypothesized to influence habitat selection by dispersers is the natal environment experienced by juveniles. Nonetheless, evidence for the effect of natal environment on dispersing, wild vertebrates remains limited. Using 18 years of nesting and mark–resight data across an entire North American geographical range of an endangered bird, the snail kite (Rostrhamus sociabilis), we tested for natal effects on breeding-site selection by dispersers and its consequences for reproductive success and population structure. Dispersing snail kites were more likely to nest in wetlands of the same habitat type (lacustrine or palustrine) as their natal wetland, independent of dispersal distance, but this preference declined with age and if individuals were born during droughts. Importantly, dispersing kites that bred in natal-like habitats had lower nest success and productivity than kites that did not. These behaviours help explain recently described population connectivity and spatial structure across their geographical range and reveal that assortative breeding is occurring, where birds are more likely to breed with individuals born in the same wetland type as their natal habitat. Natal environments can thus have long-term and large-scale effects on populations in nature, even in highly mobile animals.     

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.     

Field observations on the population dynamics of Bulinus globosus, the intermediate host of Schistosoma haematobium in the Ifakara area, Tanzania

The population dynamics of Bulinus globosus were studied in a stream and an adjacent pool near Ifakara, Tanzania. Experiments with caged snails were used to determine fecundity, hatching rates, growth and mortality. Growth and mortality rates were also estimated for the natural population from successive size frequency distributions. Oviposition began in snails as small as 6.6 mm and there was a positive correlation between the size of the snails at oviposition and the number of embryos per egg mass. Fecundity was season-dependent. In the pool, snail growth was constant throughout the year, while in the stream, growth was markedly reduced during the rainy season. Mortality rates correlated with fluctuations in snail population density and were highest during the rainy season and among young snails. The high intrinsic rate of natural increase observed for B. globosus enables it to exist in habitats which alternate frequently between favorable and adverse conditions.     

Microsatellite polymorphism in the endangered snail kite reveals a panmictic,low diversity population

Genetic structure and genetic diversity are key population characteristics that can inform conservation decisions, such as delineating management units or assessing potential risks for inbreeding depression. Evidence of genetic structuring or low genetic diversity in the critically endangered snail kite (Rostrhamus sociabilis plumbeus) would have implications for monitoring and planning decisions. Recent work on understanding connectivity across the snail kite range indicated that there is less dispersal between northern and southern parts of the current range, and that dispersal is shaped by individual habitat preference. We examine whether there is neutral genetic structure and the amount of genetic variation in the population by non-lethally sampling 235 nestlings from unique nests across the entire breeding range between 2013 and 2014. Data on 15 microsatellite revealed low diversity (e.g., N _a?=?2.54, H _e?=?0.37) and range-wide panmixia based on AMOVA, Bayesian clustering, spatial autocorrelation, isolation by distance, and spatially explicit ordination analyses. Our results emphasize that long-term recovery goals and management strategies should be based on viewing snail kites as a single genetic population, despite evidence for non-random dispersal between wetlands over ecological time scales. These results also highlight the need to understand potential effects of low genetic diversity on population dynamics and viability of snail kites. More broadly, these results add to the growing evidence for potential discrepancies between dispersal and genetic patterns, emphasizing that care should be taken if using one to interpret the other, particularly for widely-ranging species.     

Leech predation on juvenile freshwater snails: effects of size,species and substrate

Summary The leechGlossiphonia complanata does not appear to have substantial impact on snail populations, but this may be due to most studies focusing on adult snails rather than juvenile snails. In this study I investigated how predation rates ofG. complanata feeding on newly-hatched and juvenile snails was affected by snail species, snail size, snail density and substrate, in a laboratory experiment. Number of snails eaten increased with increasing density resulting in a type II functional response curve. Predation rates were higher when leeches were feeding onLymnaea emarginata than onPhysa gyrina, whereas there was no significant difference in predation rates when they were feeding onL. emarginata andHelisoma anceps. Sandy substrates and greater snail size resulted in decreased predation rates. Sand reduced movement speed ofG. complanata, which probably reduced encounter rates. Thus, there was a comparatively large effect of leech predation on newly-hatched snails, due to a high probability of encounter and high predation rates, but spatial and temporal refuges probably reduce the importance of leech predation as a structuring force in freshwater snail assemblages.     

Schistosoma mansoni: influence of Biomphalaria glabrata size on susceptibility to infection and resultant cercarial production

Biomphalaria glabrata snails of the same age, but different sizes, were used to determine size-related susceptibility to Schistosoma mansoni miracidial infection and the influence of snail size on total cercarial production. Snails with shell diameters from less than 5 to greater than 17 mm were individually exposed to one or several miracidia, depending on the experiment. In snails exposed to multiple numbers of miracidia, the percentage of snails which developed patent infections was lower in snails with larger shell sizes. This was also reflected by fewer primary sporocysts per infected snail found in tissues of the larger snails. Upon determining cercarial production in these groups over a 1-month period there were no statistical differences between any groups in the numbers of cercariae produced per snail. However, upon determining the number of successful primary sporocysts found in cohort snails of each size group, cercarial production increased as a function of the number of successful primary sporocysts. This was verified by examining cercarial production in various size snails with known monomiracidial infections. Our data therefore confirm and extend earlier work using snails infected with unknown numbers of miracidia and clearly show that total S. mansoni cercarial development and decreased susceptibility of snails is a direct reflection of snail size and not necessarily age of the snail.     

Host life history and the effect of parasitic castration on growth: A field study of Cerithidea californica Haldeman (Gastropoda : Prosobranchia) and its trematode parasites

The prevalence of parasitic infection by larval digenetic trematodes in natural populations of the mud snail, Cerithidea californica Haldeman, was found to increase with snail length; all snails ≥ 33 mm were infected. Distributions of infections by the seven most common larval trematodes were heterogeneous due to two species being more common than expected in the smaller size classes of snails, two being more common than expected in the larger size-classes of snails and three species being most prevalent in snails of intermediate length. The relative abundances of trematodes in different size-classes reflected these distributional patterns.A mark-recapture field study of snail growth rates failed to demonstrate that parasitic infection causes gigantism in Cerithidea. Parasitism tended to stunt the growth of juvenile snails and to a lesser degree, that of adult snails. The effects of trematodes on snail growth was shown to be species specific. This finding contrasts with those of earlier studies in which gigantic growth was observed in infected snails. This discrepancy is attributed to differences in the life histories of the host snails. It is predicted that gigantism will occur commonly in short-lived or semelparous species of snails but rarely, if ever, in long-lived iteroparous species which are predominately marine.     

不同施药方式对福建烟蚜抗药性及活性酶的影响

本文采用连续施用和轮换施用农药方式对福建烟区烟蚜进行抗药性及其体内酶变化的测定,研究结果表明,在连续施用3次、5次、7次和9次后,施用灭多威后的烟蚜抗性分别增长了1.397倍、2.608倍、4.891倍和7.598倍;施用吡虫啉则分别增长了1.717倍、2.114倍、2.861倍和4.169倍;施用高效氯氟氰菊酯则分别...