Morphological and life history responses to predators in larvae of seven anurans

Amphibian larvae often face two major sources of mortality: pond desiccation and predation. Tadpoles of seven anuran species with different preferences for type of breeding habitat, on a hydroperiod scale, were tested for responses to the presence of predators by raising them experimentally in the presence and absence of a separately caged invertebrate predator that was fed on conspecific tadpoles. The species typically breeding in temporary or semi-permanent ponds ( Rana temporaria , Rana arvalis , Rana dalmatina and Hyla arborea ) – where invertebrate predator populations are predicted to vary considerably spatiotemporally – all showed marked induced increases in tail fin depth in response to predator presence. These species also tended to respond by reduced growth rates. The representative of the most ephemeral habitats, Bufo calamita , did not respond in any of these traits. Its congeneric, Bufo bufo , a toxic inhabitant of permanent ponds and lakes, tended to respond to predator presence by reducing its growth rate, though not by a tail depth increase. I argue that the rather poor swimming performance in Bufo tadpoles may opt for defences other than locomotor ability. The palatable, permanent pond species Pelobates fuscus did not alter either its growth rate or tail morphology. Possible explanations for this result are discussed.     

Non-interactive multiple predator effects on tadpole survival

Interactions among and within three species of predators were estimated in terms of their effects on prey survival using short-term predation experiments. The prey were tadpoles (Rana temporaria), and the predators were dragonfly larvae (Anax imperator), newts (Triturus alpestris), and backswimmers (Notonecta glauca). Mortality rate per predator imposed by Triturus and Notonecta did not decline with predator density, whereas the predation rate of Anax was strongly reduced when the number of predator individuals increased. Impacts of all three predators were not altered by the presence of other species in pairwise combinations. This system is therefore characterized by interference between individual dragonflies but relatively independent effects of predator species. These results were largely predictable based on the natural history of the predators and are encouraging for attempts to model communities as assemblages of interacting species.     

Behavioural responses to predator chemical cues and local variation in antipredator performance in Rana temporaria tadpoles

Antipredator behaviour is an important factor influencing survival probability of prey animals, and it may evolve rapidly as a response to changes in predator regime. I studied antipredator behaviour of common frog ( Rana temporaria ) tadpoles from three populations that differ in predator regimes. In the first experiment, tadpoles obtained from four natural matings in each population were subjected to chemical cues from either European perch ( Perca fluviatilis ) or from larvae of the dragonfly Aeshna juncea . Tadpoles decreased their activity in response to both predators, but the spatial behaviour of tadpoles differed between the two predator treatments. In general, there were no differences in behaviours among the populations, but in three out of four studied behaviours there were differences between parentages within the populations suggesting that these behaviours may be genetically determined. The lack of a significant Predator×Population interaction suggests no differences in plastic antipredator behaviour among the populations, while the lack of significant Predator×Parentage interaction suggests no genetic variance within the populations for plastic antipredator behaviour. In the second experiment, tadpoles from the three populations were exposed to predation by a free-ranging A. juncea . In line with the first experiment, there were no differences in survival rate between the populations. R. temporaria tadpoles seem to rely heavily on plastic antipredator behaviour as their main response to predator chemical cues. There was very little indication of local behavioural differentiation and the possible reasons for the lack of divergence among populations are discussed.     

The effects of perceived predation risk on pre- and post-metamorphic phenotypes in the common frog

Where organisms undergo radical changes in habitat during ontogeny, dramatic phenotypic reshaping may be required. However, physiological and functional interrelationships may constrain the extent to which an individual's phenotype can be equally well adapted to their habitat throughout the life cycle. The phenotypic response of tadpoles to the presence of a predator has been reported for several species of anuran but the potential post-metamorphic consequences have rarely been considered. We reared common frog Rana temporaria tadpoles in the presence or absence of a larval odonate predator, Aeshna juncea , and examined the consequences of the resulting phenotypic adjustment in the aquatic larval stage of the life cycle for the terrestrial juvenile phenotype. In early development tadpoles developed deeper tail fins and muscles in response to the predator and, in experimental trials, swam further than those reared in the absence of a predator. While the difference in swimming ability remained significant throughout the larval period, by the onset of metamorphosis we could no longer detect any differences in the morphological parameters measured. The corresponding post-metamorphic phenotypes also did not initially differ in terms of morphology. At 12 weeks post-metamorphosis, however, froglets that developed from predator-exposed tadpoles swam more slowly and less far than those that developed from tadpoles reared in the absence of predators, the opposite trend to that observed in the larval stage of the life cycle, and had narrower femurs. These results suggest that there may be long-term costs for subsequent life-history stages of tailoring the larval phenotype to prevailing environmental conditions.     

Basal metabolic rate and risk‐taking behaviour in birds

Basal metabolic rate (BMR) constitutes the minimal metabolic rate in the zone of thermo‐neutrality, where heat production is not elevated for temperature regulation. BMR thus constitutes the minimum metabolic rate that is required for maintenance. Interspecific variation in BMR in birds is correlated with food habits, climate, habitat, flight activity, torpor, altitude, and migration, although the selective forces involved in the evolution of these presumed adaptations are not always obvious. I suggest that BMR constitutes the minimum level required for maintenance, and that variation in this minimum level reflects the fitness costs and benefits in terms of ability to respond to selective agents like predators, implying that an elevated level of BMR is a cost of wariness towards predators. This hypothesis predicts a positive relationship between BMR and measures of risk taking such as flight initiation distance (FID) of individuals approached by a potential predator. Consistent with this suggestion, I show in a comparative analysis of 76 bird species that species with higher BMR for their body mass have longer FID when approached by a potential predator. This effect was independent of potentially confounding variables and similarity among species due to common phylogenetic descent. These results imply that BMR is positively related to risk‐taking behaviour, and that predation constitutes a neglected factor in the evolution of BMR.     

Evaluation of the state of the gene pool of natural populations of vertebrate animals inhabiting the fragmented landscape of Moscow and Moscow district (with special reference to brown frogs)

A system of urban ecological genetic monitoring has been developed for the first time in Russia and in the world. As a model, two species of brown frogs, Rana arvalis Nilss. and R. temporaria L. (16 populations, seven isozyme loci) were used. An evaluation of the gene pool state in urban frog populations has shown that the diversity in the Moscow frog populations was lower than that in natural populations (up to 80 and 50% in respectively R. anvalis and R. temporaria). Mean heterozygosities per locus were higher in large natural populations than in small urban isolates: in R. arvalis, these values were 0.16 and 0.06; in R. temporaria, 0.34 and 0.18, respectively. The number of polymorphic loci was also higher in natural populations than in the urban ones: 4 versus 2 in R. arvalis and 5 versus 4 in R. temporaria. Using superoxiddismutase as an example, fixation of different alleles of the same locus in different small isolates was shown. The gene pool condition of all but one urban populations of brown frog was evaluated as unsatisfactory, and that of the R. arvalis populations, as critical. These changes of the gene pool are explained mainly by gene drift accompanied by inbreeding, which was caused by human-induced fragmentation of the range and a decrease in population size of the species. The results of this study was employed in the development of the Moscow governmental program on restoration of the gene pools of vanishing animal species on specially protected natural urban territories. The series of works on long-term monitoring and assessment of the state of natural populations of model species in anthropogenic landscapes of Moscow and Moscow district has laid a foundation for a new branch of science, gene urbanology.     

Recognition and Response to Native and Novel Predators in the Largespring mosquitofish,Gambusia geiseri

The introduction of predator species into new habitats is an increasingly common consequence of human activities, and the persistence of native prey species depends upon their response to these novel predators. In this study, we examined whether the Largespring mosquitofish, Gambusia geiseri exhibited antipredator behavior and/or an elevation of circulating stress hormones (cortisol) to visual and chemical cues from a native predator, a novel predator, or a non‐predatory control fish. Prey showed the most pronounced antipredator response to the native predator treatment, by moving away from the stimulus, while the prey showed no significant changes in their vertical or horizontal position in response to the novel or non‐predator treatments. We also found no significant difference in water‐borne cortisol release rates following any of the treatments. Our results suggest the prey did not recognize and exhibit antipredator behavior to the novel predator, and we infer that this predator species could be detrimental if it expands into the range of this prey species. Further, our study demonstrates prey may not respond to an invasive predator that is phylogenetically, behaviorally, and morphologically dissimilar from the prey species' native predators.     

Contrasting population structures in two sympatric anurans: implications for species conservation

A general prediction of the neutral theory of evolution is that genetic diversity should correlate positively with effective population size. We show here that diversity across eight microsatellite loci was consistently and substantially lower in one common amphibian (Bufo bufo) than in another with similar life history traits (Rana temporaria) despite B. bufo having the larger breeding assemblage sizes. However, B. bufo breeding assemblages were much more highly differentiated than those of R. temporaria according to both Fst and Rst estimators. These differences occurred in shared habitats across identical geographical distances. The patterns of genetic diversity and differentiation detected in these two species were probably a consequence of high gene flow in R. temporaria but much lower gene flow among the larger but more dispersed B. bufo assemblages. These observations highlight the difficulty of defining the boundaries of wild populations, and show how two broadly similar species can exhibit very different population dynamics.     

A Comparative Study of the Aminergic Structures in the Hypothalamus and the Hypophysis of Amphibian Larvae,with Special Reference to the Innervation of the Pars Distalis

The distribution and occurrence of the aminergic structures of the hypothalamus and the hypophysis of the investigated species are very similar, except for the pars distalis. In the tadpoles of the anuran species Rana temporaria, R. arvalis, R. dalmatina, R. escutenta, Bufo bufo and B. calamita, the pars distalis is aminergically innervated (the Falck-Hillarp technique), but these nerves disappear at the metamorphic climax. In the larvae of the urodelan species Triturus cristatus and T. vulgaris, the pars distalis is devoid of such nerves. Electron microscopical studies of R. temporaria, B. bufo and T. cristatus confirm these observations. A discussion of the functional significance of the pars distalis nerves and the phylogenetic role of the difference between anuran and urodelan species is given.     

Evidence for aposematism in the plant bug Lopidea nigridea Uhler (Hemiptera: Miridae: Orthotylinae)

The biology of the plant bug Lopidea nigridea is described, with particular focus on whether this red and black species can be considered aposematic. Lopidea individuals were rejected after attack in 92 of 112 feeding trials involving three of their five most common arthropod predator species. Compared to another syntopic plant bug species, L. nigridea is aggregated in distribution, and individuals are sedentary in their basic daily habits. These life-history features, coupled with the ability to survive most attacks by arthropod predators in the feeding trials, suggest that L. nigridea is protected to some degree by an aposematic defence. How L. nigridea may acquire its distastefulness is discussed, with special attention to host plant relations in L. nigridea and in the genus Lopidea as a whole.     

Feeding habits of the predaceous nymphs of Dolania Americana in Northwestern Florida (Ephemeroptera : Behningiidae)

Feeding habits of nymphs of the predatory mayflyDolania americana Edmunds & Traver in northwestern Florida were examined during 1971 and 1972. Larvae of Chironomidae (Diptera) formed the bulk of prey items. The size of the predaceousD. americana nymphs was correlated with the size of chironomid prey taken. Differential microhabit utilization allows partition of prey resources with the other major predator species present. The relationship between predator and prey biomass throughout the year is discussed.     

The pattern of lion predation in Nairobi Park

The feeding habits of lions in Nairobi National Park have been observed between 1968 and 1972. Nairobi Park is briefly described and the herbivore and predator populations are discussed. The relationship between the changing status of the prey population for the period 1968-72 and the feeding habits of the lions is examined and compared with data from earlier years. Selection ratio (preference ratio) for three of the most important prey species is more closely adjusted to the abundance of these species in the total ungulate population than was the case prior to 1968. Only warthog and eland form an exception. Predation on wildebeest shows the most marked decrease although this species has been increasing during the last 5 years. Wildebeest, kongoni, zebra and warthog have provided 80% of total kills throughout the study on the basis of numbers killed. On this basis, warthog has the highest preference rating. However, the largest contribution to the lions' diet expressed in kg is by kongoni, followed by zebra, wildebeest, eland and warthog, in that order. Kongoni and wildebeest contribute most during the first half of the year and zebra and eland during the last 6 months. Selection for males is significant in zebra and eland and highly significant in kongoni, whilst only in zebra is selection for juveniles evident.     

Behavioral diversity of predatory arboreal ants in coffee agroecosystems

Aspects of predator assemblages that alter predator effects on prey have received extensive recent attention. Among other mechanisms, differences in behavior or resource use within predator trophic levels may enhance predator effects on prey, especially if effects of each predator species differ with environmental conditions. We address whether three common ant species (Azteca instabilis F. Smith, Camponotus textor Forel, and Crematogaster spp.) are functionally unique in coffee agroecosystems, asking if each species differs in (1) cooperative foraging behavior, (2) responses to experimentally introduced herbivores, and (3) responses to pest outbreaks. Furthermore, we examined the behaviors and effects of each ant species under different conditions by varying herbivore species, herbivore size, and herbivore density and carrying out observations in different seasons. Ant species significantly differed in foraging behaviors, in effects on individual herbivores, and in responses to pest outbreaks in terms of both type and time of response to herbivores. The behaviors and effects of each ant species differed in the dry and wet seasons and for different herbivore species and sizes. Although A. instabilis generally removed more larvae and more quickly removed larvae, this was not the case under all conditions. The data presented thus support that common ant species in coffee agroecosystems are behaviorally diverse in their responses to herbivores under different conditions. We discuss the implications of these differences in ant behaviors for enhancement of predatory function in light of both multipredator effects and in terms of the potential importance of predator diversity.     

The biological function of gulls' attraction towards predators

Many species of birds and mammals are attracted towards some of their natural predators, often without overt aggression. The biological function of this predator attraction was studied in a colony of herring gulls and lesser black-backed gulls, with the aid of predator models. More birds flocked above the predator model when it was presented together with a dead gull; also the birds did not land as close to a predator with a dead gull as they did to a predator alone. Having seen a predator with a dead gull had a clear effect on the distance at which birds later avoided that particular predator on the ground when it was presented to them alone. It is suggested that predator-attraction enables animals to collect information about a potential enemy; in this information the experience of conspecifics with the predator is taken into account; subsequent behaviour of the animals depends partly on this information. The increase in the avoidance distance with respect to a predator with a dead bird may be an adaptation to carnivores' habit of surplus killing.     

Planktonic encounter rates in homogeneous isotropic turbulence: the case of predators with limited fields of sensory perception

It is a well-established fact that encounter rates between different species of planktonic microorganism, either swimming, or passively advected by the flow, are enhanced in the presence of turbulence. However, due to the complexity of the various calculations involved, current encounter rate theories are based on a number of simplifying approximations, which do not reflect reality. In particular, a typical planktonic predator is usually assumed to have perfect 'all round vision', i.e. it can perceive a prey particle at any relative orientation, provided it lies within some given contact radius R. Unfortunately, there is a wide body of experimental evidence that this is not the case. In this study the encounter problem for a predator with a limited field of sensory perception, swimming in a turbulent flow, is examined from first principles and a number of new modelling ideas proposed. A wide range of kinematic simulations are also undertaken to test these predictions. Particular attention is paid to the swimming strategy such a predator might undertake to enhance its encounter rate. It turns out that the predicted optimum swimming strategies differ radically from the results of previous work. Empirical evidence is also presented which appears to support these new findings.     

The Structure of the Mammalian Predator Guild in the Santa Cruz Formation (Late Early Miocene)

The Santa Cruz Formation (late early Miocene, Santacrucian age) registers 11 species of mammalian predators (Metatheria, Sparassodonta). Together with large carnivorous flightless birds, they comprised the terrestrial predator guild. The Santacrucian sparassodonts were diverse in body size, had different locomotory habits, and were primarily hypercarnivores. The objective of this work is to analyze the guild structure of the sparassodonts of the Santa Cruz Formation, using the variables of body mass, diet, and locomotion as proxies. Furthermore, we analyze the interaction with other predators and potential prey. The univariated test V of Poole and Rathcke and the multivariated test of Clark-Evans were used to construct the models. In the multivariate test, we made a Principal Component Analysis to resume and standardize the variables. With body mass and locomotion we obtained an evenly spaced pattern of segregation for the sparassodont species, being non-significant and significant, respectively. The pattern was aggregated and significant only with diet. The analysis of all variables together resulted in an evenly spaced and significant pattern, which is consistent with character displacements (segregation of species throughout the morphospace) that would help to diminish interspecific competition during the Santacrucian age and would allow selection of prey species of different sizes and substrate specializations. When the body size pattern of predator birds and sparassodonts were plotted together, the pattern is evenly spaced and non-significant. Other factors, including locomotion, would differentiate these species and their ecological niches.     

Diets and trophic‐guild structure of a diverse fish assemblage in Chesapeake Bay,U.S.A.

Dietary habits and trophic‐guild structure were examined in a fish assemblage (47 species) of the Chesapeake Bay estuary, U.S.A., using 10 years of data from >25 000 fish stomachs. The assemblage was comprised of 10 statistically significant trophic guilds that were principally differentiated by the relative amounts of Mysida, Bivalvia, Polychaeta, Teleostei and other Crustacea in the diets. These guilds were broadly aggregated into five trophic categories: piscivores, zooplanktivores, benthivores, crustacivores and miscellaneous consumers. Food web structure was largely dictated by gradients in habitat (benthic to pelagic) and prey size. Size classes within piscivorous species were more likely to be classified into different guilds, reflecting stronger dietary changes through ontogeny relative to benthivores and other guilds. Relative to predator species and predator size, the month of sampling had negligible effects on dietary differences within the assemblage. A majority of sampled fishes derived most of their nutrition from non‐pelagic prey sources, suggesting a strong coupling of fish production to benthic and demersal food resources. Mysida (predominantly the opossum shrimp Neomysis americana) contributed substantially to the diets of over 25% of the sampled predator groups, indicating that this species is a critical, but underappreciated, node in the Chesapeake Bay food web.     

Spatial and trophic niche differentiation in two sympatric lizards (Tropidurus torquatus and Cnemidophorus ocellifer) with different foraging tactics

Abstract In a ‘restinga’ habitat of southeastern Brazil, we studied the food habits and the microhabitat use of two lizards with distinct foraging modes: the tropidurid Tropidurus torquatus, a sit-and-wait predator, and the teiid Cnemidophorus ocellifer, a wide forager. The diet of the two species differed strongly, indicating a low level of similarity in their trophic niche. The sit-and-wait predator fed mainly on mobile prey, whereas the wide forager fed mainly on sedentary prey (larvae). The spatial niche breadth of T. torquatus was larger than that of C. ocellifer. Despite interspecific differences, the two species overlapped greatly in micro-habitat use. The data indicate that at Linhares the two lizard species differed more in food resources than in microhabitat, and that most of the food differences reflect the foraging patterns of the species.     

The schooling ethology of Lutianus monostigma (Pisces) in the shallow reef environment of Aldabra

Field observations on a school of Lutianus monostigma have been made off Aldabra in the shallow reef environment. The school does not exhibit any obvious response to temperature. The response to current is the main factor that determines school orientation and position in the channel. Longshore movements are performed but their function is not known. The morning and evening dispersal may be associated with changes in light intensity and/or the crepuscular feeding habits. The response to predators depends upon the behaviour of the predator, its species and size. Other factors affecting the school in their natural environment are described.