Parallel speciation in Astyanax cave fish (Teleostei) in Northern Mexico

We investigated differentiation processes in the Neotropical fish Astyanax that represents a model system for examining adaptation to caves, including regressive evolution. In particular, we analyzed microsatellite and mitochondrial data of seven cave and seven surface populations from Mexico to test whether the evolution of the cave fish represents a case of parallel evolution. Our data revealed that Astyanax invaded northern Mexico across the Trans-Mexican Volcanic Belt at least three times and that populations of all three invasions adapted to subterranean habitats. Significant differentiation was found between the cave and surface populations. We did not observe gene flow between the strongly eye and pigment reduced old cave populations (Sabinos, Tinaja, Pachon) and the surface fish, even when syntopically occurring like in Yerbaniz cave. Little gene flow, if any, was found between cave populations, which are variable in eye and pigmentation (Micos, Chica, Caballo Moro caves), and surface fish. This suggests that the variability is due to their more recent origin rather than to hybridization. Finally, admixture of the young Chica cave fish population with nuclear markers from older cave fish demonstrates that gene flow between populations that independently colonized caves occurs. Thus, all criteria of parallel speciation are fulfilled. Moreover, the microsatellite data provide evidence that two co-occurring groups with small sunken eyes and externally visible eyes, respectively, differentiated within the partly lightened Caballo Moro karst window cave and might represent an example for incipient sympatric speciation.     

Genetic divergence between cave and surface populations of Astyanax in Mexico (Characidae,Teleostei)

A study of genetic diversity at microsatellite loci and the mitochondrial DNA (mtDNA) cytochrome b gene was carried out to assess genetic relationships among four Mexican cave (Pachon, Sabinos, Tinaja, Chica) and four surface populations of Astyanax fasciatus (Characidae) from northeast Mexico and the Yucatan. With the exception of Chica, the cave populations were all characterized by extremely low microsatellite variability, which most likely resulted from bottleneck events. Population analyses of the microsatellite data indicated no measurable levels of gene flow between all cave and surface populations (F(ST) > 0.0707). Phylogenetic analyses of mtDNA data showed that only two cave populations - Sabinos and Tinaja - group together to the exclusion of surface populations. From the microsatellite data these cave populations cluster with the Pachon cave fish population. The mtDNA thus appears to have been replaced in Pachon because of introgressive hybridization. It is likely that these three cave populations have descended from a surface ancestor in common with current surface populations, rather than evolving recently from one of the extant surface populations. Like Pachon, the Chica population clustered with the surface populations according to mtDNA data, but was not clearly associated with either the surface or the other cave populations according to the microsatellite data. Our data indicate that the Chica population evolved recently from a surface population, and subsequently hybridized with a phylogenetically older cave population. In conclusion, both the microsatellite and mtDNA data suggest multiple origins of cave populations and the Chica and Sabinos/Tinaja/Pachon were founded after at least two independent invasions from surface populations.     

Evolutionary convergence on sleep loss in cavefish populations

Patterns of sleep vary widely among species, but the functional and evolutionary principles responsible for this diversity remain unknown. The characin fish, Astyanax mexicanus, has eyed surface and numerous blind cave populations. The cave populations are largely independent in their origins, and the species is ideal for studying the genetic bases of convergent evolution. Here we show that this system is also uniquely valuable for the investigation of variability in patterns of sleep. We find that a clearly defined change in ecological conditions, from surface to cave, is correlated with a dramatic reduction in sleep in three independently derived cave populations of A. mexicanus. Analyses of surface × cave hybrids show that the alleles for reduced sleep in the Pachón and Tinaja cave populations are dominant in effect to the surface alleles. Genetic analysis of hybrids between surface and Pachón cavefish suggests that only a small number of loci with dominant effects are involved. Our results demonstrate that sleep is an evolutionarily labile phenotype, highly responsive to changes in ecological conditions. To our knowledge, this is the first example of a single species with a convergence on sleep loss exhibited by several independently evolved populations correlated with population-specific ecologies.     

Circadian rhythms and photic entrainment of swimming activity in cave-dwelling fish Astyanax mexicanus (Actinopterygii: Characidae), from El Sotano La Tinaja,San Luis Potosi,Mexico

Circadian regulation has a profound adaptive meaning in timing the best performance of biological functions in a cyclic niche. However, in cave-dwelling animals (troglobitic), a lack of photic cyclic environment may represent a disadvantage for persistence of circadian rhythms. There are different populations of cave-dwelling fish Astyanax mexicanus in caves of the Sierra El Abra, Mexico, with different evolutive history. In the present work, we report that fish collected from El Sótano la Tinaja show circadian rhythms of swimming activity in laboratory conditions. Rhythms observed in some of the organisms entrain to either continuous light–dark cycles or discrete skeleton photoperiods tested. Our results indicate that circadian rhythm of swimming activity and their ability to entrain in discrete and continuous photoperiods persist in some organisms that might represent one of the oldest populations of cave-dwelling A. mexicanus in the Sierra El Abra.     

Evolution of an adaptive behavior and its sensory receptors promotes eye regression in blind cavefish

Background

How and why animals lose eyesight during adaptation to the dark and food-limited cave environment has puzzled biologists since the time of Darwin. More recently, several different adaptive hypotheses have been proposed to explain eye degeneration based on studies in the teleost Astyanax mexicanus, which consists of blind cave-dwelling (cavefish) and sighted surface-dwelling (surface fish) forms. One of these hypotheses is that eye regression is the result of indirect selection for constructive characters that are negatively linked to eye development through the pleiotropic effects of Sonic Hedgehog (SHH) signaling. However, subsequent genetic analyses suggested that other mechanisms also contribute to eye regression in Astyanax cavefish. Here, we introduce a new approach to this problem by investigating the phenotypic and genetic relationships between a suite of non-visual constructive traits and eye regression.

Results

Using quantitative genetic analysis of crosses between surface fish, the Pachón cavefish population and their hybrid progeny, we show that the adaptive vibration attraction behavior (VAB) and its sensory receptors, superficial neuromasts (SN) specifically found within the cavefish eye orbit (EO), are genetically correlated with reduced eye size. The quantitative trait loci (QTL) for these three traits form two clusters of congruent or overlapping QTL on Astyanax linkage groups (LG) 2 and 17, but not at the shh locus on LG 13. Ablation of EO SN in cavefish demonstrated a major role for these sensory receptors in VAB expression. Furthermore, experimental induction of eye regression in surface fish via shh overexpression showed that the absence of eyes was insufficient to promote the appearance of VAB or EO SN.

Conclusions

We conclude that natural selection for the enhancement of VAB and EO SN indirectly promotes eye regression in the Pachón cavefish population through an antagonistic relationship involving genetic linkage or pleiotropy among the genetic factors underlying these traits. This study demonstrates a trade-off between the evolution of a non-visual sensory system and eye regression during the adaptive evolution of Astyanax to the cave environment.

     

Effects of organic matter availability on the life history and production of a top vertebrate predator (Plethodontidae: Gyrinophilus palleucus) in two cave streams

1. Surface ecosystems provide the primary source of organic matter to many cave communities. Variation in the strength of connectivity to the surface suggests that some caves may be more resource‐limited than others. To test this, we examined diet, prey availability and production of an obligate cave salamander Gyrinophilus palleucus (Plethodontidae), a top predator, in two south‐eastern U.S.A. caves with different levels of organic matter (Tony Sinks cave, 165 g AFDM m^?2; Bluff River cave, 62 g AFDM m^?2). 2. We quantified density, biomass, growth rate, production and diet of G. palleucus monthly for 21 months. Diet composition, differences in prey communities and seasonal patterns in prey consumption were also analysed. 3. Salamander density, biomass and secondary production were significantly greater in the high organic matter cave (0.10 m^?2, 0.18 g AFDM m^?2, 0.12 g AFDM m^?2 year^?1) than in the low organic matter cave (0.03 m^?2, 0.03 g AFDM m^?2, 0.01 g AFDM m^?2 year^?1). Although growth rates were not statistically different between the two cave salamander populations, low recaptures probably influenced this result. 4. Isopoda prey were the major contributor to salamander production in the high organic matter cave (69%). In the low organic matter cave, production was provided by isopods (41%) and oligochaetes (20%). The lower number of prey taxa contributing to salamander production in the high organic matter cave suggests the ability to forage more selectively. 5. The differences in foraging strategy, density, biomass and secondary production were probably related to differences in the strength of surface connectivity, which controls organic matter supply. Links between basal resource level and top predator performance show the importance of bottom‐up limitation in the food webs of caves and other detritus‐based ecosystems.     

Der Micos-Fisch,Höhlenfisch in statu nascendi oder Bastard ?

The Micos Population–cave fish in statu nascendi or hybrid? Observations on the evolution of cavernicoles The Micos-Cave in the Sierra de la Colmena in the State of San Luis Potosi, Mexico, contains a cavernicolous population of Astyanax mexicanus, whose members are for the most part blind, but, in contrast to other cave dwelling populations of the same species, appear almost normal in their pigmentation. Besides these, there are also large eyed and pigmented specimens to be found in this cave. Any transitional stages between the blind and the normal visioned fish are lacking. Offspring of the blind cavernicoles that are raised under light conditions develop a superficially lying eye which is markedly smaller than normal, attaining its size proportional to the light-intensity of the experimental conditions. The size and structure of the eye-remnants of the blind fish as well as the eyes of their offspring are considerably more variable than in the river specimens. Crossings of the blind Micos fish with the river fish Astyanax and also with a blind and unpigmented troglobiont of the same species - Sabinos fish - result in both cases in a more or less intermediate F¹-hybrid. A strict inbreeding within the Micos fish, selecting specimens with especially large eyes, produced animals whose eyes are comparable to those of the river fish after only three offspring generations. Electrophoresis studies on the allozyme variability at various loci prove that the Micos fish is genetically only slightly different compared to the river fish. On the other hand in some allele frequencies there is an alternative variation between the two. The Micos fish also differs from the typical troglobionts which are monomorphic at almost all loci examined and also possess alleles that are not found in the river form. Based on the genetic constitution, the Micos fish and the river fish found in the cave do not form a panmictic population. It is also doubtful that the Micos fish is the progeny of a hybrid swarm which previously resulted from a cross between a real troglobiont and the newly arrived river fish, because the Micos fish is in every characteristic genetically very similar to the river fish, whereas no clear traces of troglobiont relationship are found. Thus the Micos fish actually appears to be a cave form in statu nascendi against which the river fish that find their way in from time to time cannot compete.     

Phylogeography of surface and cave Astyanax (Teleostei) from Central and North America based on cytochrome b sequence data

Astyanax fasciatus has become a model organism for the study of regressive and adaptive evolution in cave animals. To fully understand these processes, it is important to have background information on the systematics and phylogeography of surface and cave populations of this species. Here we investigate the phylogeography of A. fasciatus in North and Central America and also the historical biogeography of this region. Phylogenetic analysis of part of the mtDNA cytochrome b gene from 26 surface and nine cave A. fasciatus populations revealed seven major clades, which, in principle, represent geographical patterns of distribution. However, the four strongly eye and pigment reduced cave populations, Piedras, Sabinos, Tinaja, and Curva, form a separate cluster, which is not sister group to the surface populations from the same locality. Similarly the Belizean populations do not cluster with their geographic neighbors from the Yucatan. The analyses indicate that there have been recurrent invasions of surface Astyanax from the south, that were most likely influenced by major climate changes during the Pleistocene. During this period, ancestors of the strongly eye and pigment reduced cave populations were able to survive underground as thermophilic relics when the surface populations became extinct. The high level of genetic divergence among the different clades shows that differing haplotype lineages must have reinvaded the surface waters from the south and/or back-colonized them from residual habitats and also penetrated into the caves. Nested clade analyses show that recurrent gene flow as well as historic processes like past fragmentation and range expansion have influenced current populations of A. fasciatus in Central and North America. Different haplotype clades of the phylogeny are not compatible with the present taxonomy of Astyanax and, therefore, we propose the application of a single systematic unit, called A. fasciatus.     

Predation of a cave fish (Poecilia mexicana, Poeciliidae) by a giant water-bug (Belostoma, Belostomatidae) in a Mexican sulphur cave

Abstract.  1. Caves are often assumed to be predator-free environments for cave fishes. This has been proposed to be a potential benefit of colonising these otherwise harsh environments. In order to test this hypothesis, the predator–prey interaction of a belostomatid (predator) and a cave fish (prey) occurring in the Cueva del Azufre (Tabasco, Mexico) was investigated with two separate experiments.
2. In one experiment, individual Belostoma were given a chance to prey on a cave fish, the cave form of the Atlantic molly ( Poecilia mexicana ), to estimate feeding rates and size-specific prey preferences of the predator. In the other experiment, population density of Belostoma was estimated using a mark–recapture analysis in one of the cave chambers.
3. Belostomatids were found to heavily prey on cave mollies and to exhibit a prey preference for large fish. The mark–recapture analysis revealed a high population density of the heteropterans in the cave.
4. The absence of predators in caves is not a general habitat feature for cave fishes. None the less predation regimes differ strikingly between epigean and hypogean habitats. The prey preference of Belostoma indicates that cave-dwelling P. mexicana experience size-specific predation pressure comparable with surface populations, which may have implications for life-history evolution in this cave fish.     

Asymmetric competition between distant taxa: poecilid fishes and water striders

Summary Competition for water surface prey between fish (Priapichtus annectens: Poeciliidae) and water striders (Potamobates unidentatus: Gerridae) was studied in the laboratory and in pools in a small tropical stream. Laboratory experiments showed that fish depressed activity and foraging success of water striders. Large fish (4–5 cm) had a greater effect than small fish (2–3 cm). The field experiment showed that competition was highly asymmetric. Presence of fish decreased water strider foraging success while the reverse interaction was insignificant. It is suggested that the higher individual foraging success of the fish, harassment of water striders by fish and the use of an exclusive resource, benthic invertebrates, by the fish, contribute to this pattern. Habitat use differed between the two species. Fish used the deeper parts of stream pools and water striders used the shallower parts of the pools. Asymmetric interference and exploitation competition may force water striders to use shallow edge habitats.     

Does a long‐term shift in Wood Stork diet foreshadow adaptability to human‐induced rapid environmental change?

To preserve biodiversity, identifying at‐risk populations and developing conservation plans to mitigate the effects of human‐induced rapid environmental change (HIREC) are essential. Changes in diet, especially for food‐limited species, can aid in detecting populations being impacted by HIREC, and characterizing the quality, abundance, and temporal and spatial consistency of newly consumed food items may provide insight concerning the likelihood of a species persisting in a changing environment. We used Wood Storks (Mycteria americana) nesting in the Florida Everglades as a model system to study the possible effects of HIREC on a food‐limited population. We compared the diets of Wood Storks in 2013 and 2014 with those reported during the 1970s before major anthropogenic activities affected the Everglades system and prey availability. Wood Storks in our study consumed more large‐bodied sunfish species (Lepomis spp.), fewer native marsh fishes, and more non‐native fish species than during the 1970s. Large sunfish and non‐native fish are relatively rare in the drying pools of Everglades marshes where storks traditionally forage, suggesting that Wood Storks may be using novel foraging habitats such as created wetlands (i.e., canals and stormwater ponds). Although created wetlands have long hydroperiods conducive to maintaining large‐bodied fishes and could provide alternative foraging habitat when prey availability is reduced in natural marshes, additional studies are needed to determine the extent to which these wetlands are used by Wood Storks and, importantly, the quality of prey items potentially available to foraging Wood Storks in created wetlands.     

Parental genetic effects in a cavefish adaptive behavior explain disparity between nuclear and mitochondrial DNA

Epigenetic parental genetic effects are important in many biological processes but their roles in the evolution of adaptive traits and their consequences in naturally evolving populations remain to be addressed. By comparing two divergent blind cave-dwelling cavefish populations with a sighted surface-dwelling population (surface fish) of the teleost Astyanax mexicanus, we report here that convergences in vibration attraction behavior (VAB), the lateral line sensory receptors underlying this behavior, and the feeding benefits of this behavior are controlled by parental genetic effects, either maternal or paternal inheritance. From behavioral studies and mathematical evolutionary simulations, we further demonstrate that disparity in nuclear and mitochondrial DNA in one of these cavefish populations that has hybridized with surface fish can be explained by paternal inheritance of VAB. The results suggest that parental genetic effects in adaptive behaviors may be important factors in biasing mitochondrial DNA inheritance in natural populations that are subject to introgression.     

Phenotypic plasticity in the greater mouse-eared bat in extremely different roost conditions

Bats use various roost types with a wide spectrum of ecological features. The greater mouse-eared bat Myotis myotis (Borkhausen, 1797), creates nurseries in attics and caves in Central Europe. The stable low temperature and high humidity cave microclimate contrasts that of attics, which may alter species adaptations and life strategies. We analysed population characteristics (composition, body condition, parasite load, and immune response) and genetic relatedness of two proximal M. myotis populations. Age, sexual and parasite species composition were similar between the cave and attic sites. However, a significantly higher parasite load and body condition was detected in the post-partum females and juveniles of the cave colony (n = 263 bats from the cave, 231 from the attic), with the cave colony females having a significantly stronger immune response (n = 2 caves and 2 attics, 20 females per site). There was no evidence for genetic divergence between cave and attic populations (n = 3 caves and 3 attics, 24 females per site), indicating that different population characteristics are not genetically based and that M. myotis is an example of a species with rather unique phenotypic plasticity.     

Effects of spatial scale and foraging efficiency on the predictions made by spatially-explicit models of fish growth rate potential

Synopsis Spatially-explicit modeling of fish growth rate potential is a relatively new approach that uses physical and biological properties of aquatic habitats to map spatial patterns of fish growth rate potential. Recent applications of spatially-explicit models have used an arbitrary spatial scale and have assumed a fixed foraging efficiency. We evaluated the effects of spatial scale, predator foraging efficiency (combined probabilities of prey recognition, attack, capture, and ingestion), and predator spatial distribution on estimates of mean growth rate potential of chinook salmon,Oncorhynchus tshawytscha. We used actual data on prey densities and water temperatures taken from Lake Ontario during the summer, as well as, simulated data assuming binomial distribution of prey. Results show that a predator can compensate for low foraging efficiency by inhabiting the most profitable environments (regions of high growth rate potential). Differences exist in predictions of growth rate potential across spatial scales of observation and a single scale may not be adequate for interpreting model results across seasons. Continued refinements of this modeling approach must focus on the assumptions of stationary distributions of predator and prey populations and predator foraging tactics.     

Evidence for multiple genetic forms with similar eyeless phenotypes in the blind cavefish, Astyanax mexicanus

A diverse group of animals has adapted to caves and lost their eyes and pigmentation, but little is known about how these animals and their striking phenotypes have evolved. The teleost Astyanax mexicanus consists of an eyed epigean form (surface fish) and at least 29 different populations of eyeless hypogean forms (cavefish). Current alternative hypotheses suggest that adaptation to cave environments may have occurred either once or multiple times during the evolutionary history of this species. If the latter is true, the unique phenotypes of different cave-dwelling populations may result from convergence of form, and different genetic changes and developmental processes may have similar morphological consequences. Here we report an analysis of variation in the mitochondrial NADH dehydrogenase 2 (ND2) gene among different surface fish and cavefish populations. The results identify a minimum of two genetically distinctive cavefish lineages with similar eyeless phenotypes. The distinction between these divergent forms is supported by differences in the number of rib-bearing thoracic vertebrae in their axial skeletons. The geographic distribution of ND2 haplotypes is consistent with roles for multiple founder events and introgressive hybridization in the evolution of cave-related phenotypes. The existence of multiple genetic lineages makes A. mexicanus an excellent model to study convergence and the genes and developmental pathways involved in the evolution of the eye and pigment degeneration.     

Habitat overlap between predatory benthic fish and their invertebrate prey in streams: the relative influence of spatial and temporal factors on predation risk

1. The spatial heterogeneity of ecosystems as well as temporal activity patterns of organisms can have far‐reaching effects on predator–prey relationships. We hypothesised that spatiotemporal constraints in mesohabitat use by benthic fish predators would reduce habitat overlap with benthic invertebrates and lead to mesohabitat‐specific predation risks. 2. We analysed the spatiotemporal activity patterns of two small‐bodied benthivorous fishes, gudgeon (Gobio gobio) and stone loach (Barbatula barbatula), and of benthic invertebrates in a small temperate stream during three 24‐h field experiments. By applying a novel method of field video observation, we monitored the spatiotemporal foraging behaviour of the fish in their natural environment. A parallel analysis of invertebrate mesohabitat use by means of small area Hess sampling allowed a direct estimation of habitat overlap at a pool–riffle scale. 3. Gudgeon showed a dominant spatial activity pattern preferring pools at all times of day, whereas stone loach used both mesohabitats but with a distinct temporal (nocturnal) activity pattern. The patterns of residence were not identical with those of active foraging. Invertebrate community composition differed significantly between mesohabitats but not between times of day. More than half of the total dissimilarity between pools and riffles was accounted for by six invertebrate taxa. Five of these were subject to higher fish predation in pools than in riffles. The total prey consumption of the two fish species together in pools was about three times as high as in riffles. Trophic niche breadth of stone loach and thus its predation range was broader than that of gudgeon. 4. These results indicate that the potential predation risk for stream invertebrates depends on the combination of spatial and temporal patterns of both predator and prey. Given the distinct differences in predation risk found between pools and riffles, we conclude that spatial heterogeneity at the mesohabitat scale can influence mechanisms and consequences of selective predation. We also suggest that the analysis of spatiotemporal predator–prey relationships should not be based on the premise that the main residence habitat and active foraging habitat of a predator are identical.     

Microchemical provenancing of prey remains in cormorant pellets reveals the use of diverse foraging grounds

Piscivorous birds in aquatic ecosystems exert predation pressure on fish populations. But the site-specific impact on fish populations, including stocked and commercially used fish species, remains disputed. One of the key questions for the management of piscivorous birds and fish is determining the origin of prey and thus which fish populations are targeted by the birds. We addressed this question by provenancing otoliths (earstones) of fish obtained from regurgitated pellets of piscivorous birds by otolith microchemistry analysis. We retrieved otoliths from regurgitated pellets of great cormorants (Phalacrocorax carbo sinensis) collected every 2 weeks for 2 years from breeding and roosting colonies at Chiemsee in Bavaria, Germany, and classified them according to family or species. We collected water samples from Chiemsee and potential surrounding foraging grounds. We measured the strontium (Sr) ⁸⁷Sr/⁸⁶Sr isotope ratio and Sr mass fraction of water and otoliths using (laser ablation) inductively coupled plasma-mass spectrometry. We assigned otoliths from regurgitated pellets to habitat clusters of origin by comparing the Sr isotopic and elemental composition of otoliths and waterbodies. In 36% of cormorant pellets collected at Chiemsee, prey was assigned to waterbodies distinct from Chiemsee. Furthermore, cormorants used different foraging sites during 1 day. Microchemical provenancing of prey remains can contribute to identifying foraging sites of piscivorous birds and to what extend the birds switched among foraging sites.     

Milyeringa Veritas (Eleotridae), a remarkably versatile Cave Fish from the Arid Tropics of Northwestern Australia

The blind cave gudgeon Milyeringa veritas is restricted to groundwaters of Cape Range and Barrow Island, northwestern Australia. It occurs in freshwater caves and in seawater in anchialine systems. It is associated with the only other stygobitic cave vertebrate in Australia, the blind cave eel, Ophisternon candidum, the world's longest cave fish, and a diverse stygofauna comprising lineages with tethyan tracks and widely disjunct distributions, often from North Atlantic caves. The cave gudgeon inhabits a karst wetland developed in Miocene limestones in an arid area. There is an almost complete lack of information on the basic biology of this cave fish, despite it being listed as threatened under the Western Australian Wildlife Conservation Act. Allozyme frequencies and distributions indicate significant population sub-structuring on the Cape Range peninsula such that the populations are essentially isolated genetically suggesting that more than one biological species is present. Further, they suggest that the vicariant events may have been associated with a series of eustatic low sealevels. Analysis of intestinal contents indicates that they are opportunistic feeders, preying on stygofauna and accidentals trapped in the water, at least at the sites sampled which were open to the surface, a conclusion supported by the results of stable isotope ratio analysis. The gudgeons are found in freshwater caves and throughout deep anchialine systems in which they occur in vertically stratified water columns in which there is a polymodal distribution of water chemistries (temperature, pH, salinity, dissolved oxygen, redox, dissolved inorganic nitrogen series, hydrogen sulphide).     

ARE PARALLEL MORPHOLOGIES OF CAVE ORGANISMS THE RESULT OF SIMILAR SELECTION PRESSURES?

The amphipod Gammarus minus is present in both caves and springs, with cave populations showing elaborated (size and antennae) and reduced (eye) characters relative to spring populations. Earlier studies have shown that cave populations resulted from independent invasions of hydrologically isolated subterranean drainages and that there is genetic variation for both elaborated and reduced characters. In this study we tested the hypothesis that a similar pattern of selection on isolated cave populations is responsible for the parallel evolution of cave morphologies. We used variation in mating success and fecundity to test for the presence of directional selection on eye, antennal, and body size characters in a set of cave and spring populations during a series of seasonal cross-sectional samplings. We found significant directional selection for smaller eyes in caves and larger eyes in springs, which supports the hypothesis that selection is responsible for reduced eye size in cave populations. We also found selection for larger body and antennal size in cave populations, which is consistent with the hypothesis that parallel patterns of selection in caves are responsible for the elaboration of body and antennal size. However, we found selection for larger body and antennal size in spring populations that is not consistent with the observed divergence of spring and cave populations. We suggest that unmeasured components of viability selection could be more important in springs than in caves and may act against the selection for larger size found in spring populations.     

How does the presence of a conspecific individual change the behavioral game that a predator plays with its prey?

Behavioral games predators play among themselves may have profound effects on behavioral games predators play with their prey. We studied the behavioral game between predators and prey within the framework of social foraging among predators. We tested how conspecific interactions among predators (little egret) change the predator–prey behavioral game and foraging success. To do so, we examined foraging behavior of egrets alone and in pairs (male and female) in a specially designed aviary consisting of three equally spaced pools with identical initial prey (comet goldfish) densities. Each pool was comprised of a risky microhabitat, rich with food, and a safe microhabitat with no food, forcing the fish to trade off food and safety. When faced with two versus one egret, we found that fish significantly reduced activity in the risky habitat. Egrets in pairs suffered reduced foraging success (negative intraspecific density dependence) and responded to fish behavior and to their conspecific by changing their visiting regime at the different pools—having shorter, more frequent visits. The time egret spent on each visit allowed them to match their long-term capture success rate across the environment to their capture success rate in the pool, which satisfies one aspect of optimality. Overall, egrets in pairs allocated more time for foraging and changed their foraging tactics to focus more on fish under cover and fish ‘peeping’ out from their shelter. These results suggest that both prey and predator show behavioral flexibility and can adjust to changing conditions as needed in this foraging game.