Differentiated phenotypic plasticity in larvae of the cannibalistic salamander Hynobius retardatus

Alternative phenotypes in natural populations can arise from either genetic polymorphism or an environmentally induced phenotype, that is, polyphenism. Evolutionary models of polyphenism developed by theoretical studies predict that polyphenism is favored when there are environment-dependent fitness trade-offs between alternatives and that the threshold frequency for a facultative switch between alternative phenotypes is adjusted in accordance with different selection regimes. The broad-headed (alternative) larval morph of Hynobius retardatus, which is induced by crowding with conspecifics or heterospecific anuran (Rana pirica) larvae, is a representative example of cannibalistic polyphenism. Morph induction by such proximate factors must reflect evolutionary (conditional frequency-dependent) processes. To clarify the role of frequency-dependent processes in polyphenism, I investigated the occurrence rate of the broad-headed morph under experimental crowding conditions (low conspecific, high conspecific, and high heterospecific densities) using larvae from eight natural populations with different larval densities of conspecifics and heterospecifics, and found interpopulational differences in the expression of the morph. Thus, there is a larval density-dependent equilibrium frequency of the morph in each pond, suggesting that the local switch point for morph induction was modified by selection to produce evolved differences between ponds. The evolution of such interpond differences has three necessary conditions: (1) There are pond-dependent fitness trade-offs between alternatives, (2) The maintenance of the morph is costly, and (3) The presence of conspecific or, especially, heterospecific larvae provides a reliable cue to the receiver

Differentiated egg size of the cannibalistic salamander Hynobius retardatus

Larvae of the salamander, Hynobius retardatus, are carnivorous, and even though there are two morphs, a typical morph and a broad-headed or “cannibal” morph, both are cannibalistic. They also sometimes eat other large prey, for example larvae of the frog, Rana pirica. In natural habitats, use of both conspecific and R. pirica larvae as food may contribute more strongly to high survival and substantially to fitness when larval densities are higher, because early-stage H. retardatus larvae sometimes experience scarcity of their typical prey. In cannibalistic oviparous amphibians, larger individuals that developed from larger eggs can more efficiently catch and consume larger prey and thus their survival may be better than that of smaller individuals developed from smaller eggs. Populations might therefore diverge in respect of egg size in response to variation in the density of conspecific and R. pirica larvae in natural ponds, with eggs being larger when larval density is higher. I examined how variance in hatchling size correlated with the incidence of cannibalism, and whether increasing larval density in natural ponds correlated with increasing egg size. Variance in initial larval body size facilitated cannibalism, and egg size increased as larval density in the ponds increased. In ponds with high larval density, where cannibalism and large prey consumption is a critical factor in offspring fitness, the production of fewer clutches with larger eggs, and thus of fewer and larger offspring, results in greater maternal fitness. Variation among the mean egg size in populations is likely to represent a shift in optimum egg size across larval density gradients

Plasticity in the timing of a major life-history transition and resulting changes in the age structure of populations of the salamander Hynobius retardatus

Variation in age and size at life-history transitions is a reflection of the diversifying influence of biotic or abiotic environmental change. Examples abound, but it is not well understood how such environmental changes influence the age structure of a population. I experimentally investigated the effects of water temperature and food type on age and body size at metamorphosis in larvae of the salamander Hynobius retardatus. In individuals grown at a cold temperature (15 °C) or given Chironomidae as prey, the time to metamorphosis was significantly prolonged, and body size at metamorphosis was significantly enlarged, compared with individuals grown at a warmer temperature (20 °C) or fed larvae. I also examined whether larval density (a possible indicator of cannibalism in natural habitats) generated variation in the age structure of natural populations in Hokkaido, Japan, where the climate is subarctic. Natural ponds in Hokkaido may contain larvae that have overwintered for 1 or 2 years, as well as larvae of the current year, and I found that the number of age classes was related to larval density. Although cool water temperatures prolong the larval period and induce later metamorphosis, in natural ponds diet-based enhancement of development translated into a shorter larval duration and earlier metamorphosis. Geographic variation in the frequency of cannibalism resulted in population differences in metamorphic timing in H. retardatus larvae. It is important to understand how environmental effects are ultimately transduced through individual organisms into population-level phenomena, with the population response arising as the summation of individual responses. Without a thorough comprehension of the mechanisms through which population and individual responses to environmental conditions are mediated, we cannot interpret the relationship between population-level and individual-level phenomena. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102, 100–114

Plastic Responses to Different Types of Cue : Predator-Induced and Deep-Water-Induced Polyphenisms in a Salamander Hynobius retardatus

Certain plastic morphological responses of animals induced across a range of environmental conditions may be adapted for effective locomotor performance. Larvae of the salamander, Hynobius retardatus, occasionally swim upward to the surface to breathe air because aquatic respiration alone is insufficient to meet their increasing respiratory requirements for growth. We hypothesized that H. retardatus larvae living in deep water would show an induced plastic response affecting locomotor structures, namely, a deeper tail, similar to that induced by predatory dragonfly larvae (Aeschna juncea), to improve their swimming performance. In this study, larval salamanders responded similarly to different cues (waterborne chemicals in a predatory environment and distance to the water's surface) by developing deeper tails. The similar modifications in tail shape presumably increase a larva's swimming performance, thereby improving its ability both to escape an attacking predator and to swim to the surface for air. The response in tail shape induced by the predatory environment was rapid, but was more gradual in larvae raised in deep water, suggesting that animals may quickly assess a dangerous environment and immediately respond, whereas assessment of an environment not requiring an immediate response for survival may be slower, accounting for the delayed response

Likelihood Inference for Copula Models Based on Left-Truncated and Competing Risks Data from Field Studies

Survival and reliability analyses deal with incomplete failure time data, such as censored and truncated data. Recently, the classical left-truncation scheme was generalized to analyze “field data”, defined as samples collected within a fixed period. However, existing competing risks models dealing with left-truncated field data are not flexible enough. We propose copula-based competing risks models for latent failure times, permitting a flexible parametric form. We formulate maximum likelihood estimation methods under the Weibull, lognormal, and gamma distributions for the latent failure times. We conduct simulations to check the performance of the proposed methods. We finally give a real data example. We provide the R code to reproduce the simulations and data analysis results