Hynobius Retardatus Research Articles

Growth-Promotion Effects of Dissolved Amino Acids in Three Species of Hynobius Salamander Hatchlings.

It has been suggested that aquatic vertebrates may be able to meet their energy requirements by using the amino acids dissolved in environmental water. If this ability can be applied to aquatic organisms generally, then conventional ecological theories related to food web interactions should be revisited, as this would likely bring about significant advances in applications. Here, we prepared two 1 mM amino acid (phenylalanine and glycine) solutions in environmental water and conducted laboratory experiments to demonstrate the utilization of dissolved amino acids by hatchlings of three salamander species (Ezo: Hynobius retardatus, Tohoku: Hynobius lichenatus, and Japanese black: Hynobius nigrescens). Compared to controls (no amino acids in environmental water), the growth rate for Ezo salamanders was higher when larvae were reared in phenylalanine solution, while that for Japanese black salamanders was higher in glycine and phenylalanine solutions. Amino acids in environmental water had no effect on the growth of Tohoku salamanders. However, when growth was divided into early (days 1 to 5) and late (days 5 to 7) developmental stages, growth in early-developmental stage individuals was improved by phenylalanine treatment, even in Tohoku salamanders. The results showed that the growth of salamander larvae was improved when salamanders were reared in environmental water with high amino acid concentrations. Although aquatic bacteria may not have been removed completely from the environmental water, no other eukaryotes were present. Our results suggest an overlooked nutrient pathway in which aquatic vertebrates take up dissolved amino acids without mediation by other eukaryotes.

Predator cannibalism can shift prey community composition toward dominance by small prey species.

Cannibalism among predators is a key intraspecific interaction affecting their density and foraging behavior, eventually modifying the strength of predation on heterospecific prey. Interestingly, previous studies showed that cannibalism among predators can increase or reduce predation on heterospecific prey; however, we know less about the factors that lead to these outcomes. Using a simple pond community consisting of Hynobius retardatus salamander larvae and their associated prey, I report empirical evidence that cannibalism among predators can increase predation on large heterospecific prey but reduce that on small heterospecific prey. In a field‐enclosure experiment in which I manipulated the occurrence of salamander cannibalism, I found that salamander cannibalism increased predation on frog tadpoles but reduced that on aquatic insects simultaneously. The contrasting effects are most likely to be explained by prey body size. In the study system, frog tadpoles were too large for non‐cannibal salamanders to consume, while aquatic insects were within the non‐cannibals’ consumable prey size range. However, when cannibalism occurred, a few individuals that succeeded in cannibalizing reached large enough size to consume frog tadpoles. Consequently, although cannibalism among salamanders reduced their density, salamander cannibalism increased predation on large prey frog tadpoles. Meanwhile, salamander cannibalism reduced predation on small prey aquatic insects probably because of a density reduction of non‐cannibals primarily consuming aquatic insects. Body size is often correlated with various ecological traits, for instance, diet width, consumption, and excretion rates, and is thus considered a good indicator of species’ effects on ecosystem function. All this considered, cannibalism among predators could eventually affect ecosystem function by shifting the size composition of the prey community.

Paedomorphosis in the Ezo salamander (Hynobius retardatus) rediscovered after almost 90 years

Although paedomorphosis is widespread across salamander families, only two species have ever been documented to exhibit paedomorphosis in Hynobiidae. One of these two exceptional species is Hynobius retardatus in which paedomorphosis was first reported in 1924, in specimens from Lake Kuttara in Hokkaido. This population became extinct after the last observation in 1932; since then, no paedomorphs of this species have been reported anywhere. Here, we report the rediscovery of paedomorphs of this species. Three paedomorph-like male salamanders were collected from a pond in the south Hokkaido in December 2020 and April 2021; in size, these specimens were similar to metamorphosed adults but they still displayed larval features such as external gills and a well-developed caudal fin. An artificial fertilization experiment demonstrated that they were sexually compatible with metamorphosed females, thus, confirming them to be paedomorphs. Future efforts to find additional paedomorphs in this and other populations are required to assess the prevalence of paedomorphosis in H. retardatus and to improve understanding of the ecology and evolution of paedomorphisis in Urodela.

Phenotypic Plasticity of Salamander Hatchlings in the Pre-Feeding Stage in Response to Future Prey.

Vulnerability of animals immediately after hatching may induce plasticity in early ontology that becomes important for subsequent survival and growth. Ezo salamanders (Hynobius retardatus) are amphibians inhabiting ponds in Hokkaido, Japan where ezo brown frogs (Rana pirica) spawn on occasion. The salamander larvae must achieve sufficient size in order to successfully capture frog tadpoles, and we examined whether the presence of tadpoles causes development of greater body and/or gape size in newly hatched salamander larvae, which will in turn result in advantageous future prey-predator interactions. To examine this hypothesis, we conducted three laboratory experiments to demonstrate the phenotypic plasticity of salamander hatchlings in response to the presence or absence of frog tadpoles and to screen the type of signals involved in the expression of the phenotypic plasticity. First, salamander hatchlings were reared alone or with tadpoles, and the growth and morphological traits of the hatchlings were compared. The results showed that hatchling larvae grew faster with a more developed gape in the presence of tadpoles. Next, to identify the type of signals inducing this plasticity, two separate experiments with manipulated chemical and visual signals from tadpoles were conducted. The findings showed that faster growth and a more developed gape were induced by chemical but not visual signals. This plasticity may be an adaptive strategy because it increases the likelihood of preying on tadpoles in future prey-predator interactions.

Predation impacts of invasive raccoons on rare native species

In Japan, there are concerns that invasive alien raccoons prey on rare native species during their spawning season from late winter to early summer. We investigated raccoon predation impact by examining the predation presence using DNA metabarcoding and extent of predation on rare native species using stable isotope analysis. We captured raccoons in Hokkaido, Japan from April to August in 2018 and 2019. We analysed raccoon faeces and gastric contents by DNA metabarcoding to detect the rare native Hokkaido salamander and Japanese crayfish. Hokkaido salamanders were detected from gastric contents, but Japanese crayfish were not detected in any samples. Stable isotope analysis of raccoon muscle samples and the Bayesian mixing model were used to estimate each food resource’s contribution to the raccoon diet. Animal food resources accounted for 70% of total consumed food. The foraging ratios of amphibians and crustaceans were about 9% and 5%, respectively. Raccoons have been found to use amphibians at a higher rate than previously reported, including a rare endangered species, the Hokkaido salamander. Hokkaido salamander and Japanese crayfish spawn in the spring, and increased predation pressure by raccoons may directly impact populations of these rare native species.

Expression of Genes Involved in Offensive and Defensive Phenotype Induction in the Pituitary Gland of the Hokkaido Salamander (Hynobius retardatus).

Amphibians exhibit phenotypic plasticity, which allows flexible adaptation to fluctuating environments. Although genes involved in expression of plastic phenotypes have been identified, the endocrine bases of plastic responses are largely unknown. Larvae of the Hokkaido salamander (Hynobius retardatus) plastically display distinct phenotypes, an "offensive phenotype" characterized as larger body with broadened gape and a "defensive phenotype" characterized as enlarged gills and tail and less active behavior, in the presence of prey larval amphibians and predatory larval dragonfly, respectively. In the presence of both prey and predators, the degree of induction of both phenotypes is reduced, suggesting cross-talk between the molecular signaling pathways of these phenotypes. We conducted a transcriptomic analysis to examine how endocrine regulation affects the phenotypic expression by focusing on the pituitary gland. We found that five endocrine genes, i.e., calcitonin related polypeptide alpha (CALCA), growth hormone (GH), neuropeptide B (NPB), parathyroid hormone 2 (PTH2), and prolactin 1 (PRL1), were involved in the expression of both phenotypes. However, we conducted only RNA-seq analysis, and no confirmation of significant up-regulation or down-regulation has been conducted. These results suggest that these genes were up-regulated for induction of the offensive phenotype and down-regulated for induction of the defensive phenotype. Phylogenetic analysis indicated that possible gene duplications of PRL and CALCA have occurred during amphibian evolution. Based on these findings, it is suggested that a trade-off of molecular signaling pathways exists between the two distinct phenotypic expressions. The results also suggest that hormonal-gene duplications might have contributed to the acquisition of phenotypic plasticity in amphibians.

Enhanced recruitment of larger predators in the presence of large prey.

Most carnivores undergo diet shift from smaller to larger prey items during ontogeny. The trophic relationship between a growing carnivore and larger prey is representative of a size-structured predator-prey interaction. The strength of this interaction is, in part, determined by the recruitment of individuals from smaller predatory size classes into larger predatory size classes. Therefore, it is interesting to investigate how larger prey alter the recruitment of smaller predator size classes into larger predator size classes, since this can affect their own future predation risk. Past empirical studies have exclusively documented that large prey reduce predator recruitment by decreasing growth and/or survival of the smaller predators. In this study, we provide empirical evidence of the contrasting pattern: large prey enhance the recruitment of smaller predators into larger predators even though they increase cannibalism mortality of the smaller predators. We have done this here by studying the trophic interaction between predatory salamander larvae Hynobius retardatus and the frog tadpoles Rana pirica that represent their large prey. In a field experiment in which salamander hatchlings were exposed to the presence or absence of large frog tadpoles, we found that more giant salamanders emerged in the presence of frog tadpoles than in their absence. Reassignment of frog tadpoles (to both treatments) in the subsequent experimental period showed that the enhanced emergence of giant salamanders in the presence of frog tadpoles leads to the intensification of salamander predation on the frog tadpoles. In an additional laboratory experiment, to better understand the underlying mechanisms, we manipulated both the presence of frog tadpoles and the occurrence of cannibalism between salamander hatchlings. This experiment revealed that frog tadpoles intensify the cannibalism of salamander larvae during their hatchling stage, thus allowing more salamander larvae to become large-sized predators. Our results suggest that frog tadpoles can inadvertently intensify their own future predation risk by intensifying cannibalistic interactions among predatory salamander hatchlings, thereby enhancing the degree of predator recruitment to a larger size class. Hence, large prey can enhance the recruitment of individuals from small predator size classes into larger predator size classes.

Comparison of susceptibility to a toxic alien toad (Bufo japonicus formosus) between predators in its native and invaded ranges

Abstract To manage biological invasions effectively, the impacts of alien species on the demography and traits of native species must be known, but determining those impacts can be challenging. We used a comparative approach to gain insight into the impacts that an alien toad (Bufo japonicus formosus) might have on native Japanese predatory amphibians. We compared the susceptibility of native predator species to alien toad toxins in the alien‐invaded range and the susceptibility of closely related native predator species to the toxins in the alien toad's native range to investigate the impacts of an alien on a native species. Bufo japonicus formosus is native to Honshu, but was recently introduced to Hokkaido and Sado. In laboratory experiments, we compared individual mortality of predators exposed to a toad hatchling between novel predators on the toad‐invaded islands and ecologically similar congeneric or conspecific species on Honshu, where the toad is native. We also compared (1) the percentage of individuals that consumed a toad hatchling and (2) toxin resistance (i.e. survival and growth of individuals after toad consumption) between these two groups of predators, as mechanistic components behind the susceptibility of the predators to the toxic prey. The mortality of Rana pirica from all populations after consumption of a toad hatchling was almost 100%, and that of Hynobius retardatus ranged from 14 to 90%, depending on the population. In contrast, the mortality of Rana ornativentris and Hynobius nigrescens was near 0% regardless of population. These differences between congeneric predators were mostly due to differences in their toxin resistance. These results suggest that the alien toad is a potential threat to the novel amphibian predators on Hokkaido, although they also imply that the novel predators on Hokkaido have the potential to develop toxin resistance through adaptive evolution. However, this counteradaptation may have a higher chance of evolving in H. retardatus than in R. pirica because of differences in their genetic backgrounds.

Foraging traits of native predators determine their vulnerability to a toxic alien prey

Abstract Vulnerability to toxic alien prey varies among native predator species, depending on their toxin resistance. Although foraging traits relating to the intake of prey are also likely to influence predator vulnerability to toxic alien prey, these traits have received little attention. To address this issue, we studied the toxic impacts of an alien prey species (Bufo japonicus formosus toad hatchlings) on a carnivorous and an omnivorous native amphibian species (Hynobius retardatus salamander larvae and Rana pirica frog tadpoles, respectively) that both feed on alien B. japonicus formosus toad hatchlings. First, we designed a mesocosm and an aquarium experiment to evaluate the impact of B. japonicus formosus on the survival of native amphibians under various alien prey densities and predation regimes. Second, we performed several additional laboratory experiments to clarify specific processes of native amphibian mortality in the presence of the alien B. japonicus formosus toad hatchlings. We found that both native amphibian species were impacted by the alien B. japonicus formosus toad hatchlings. Interestingly, R. pirica frog tadpoles were more affected than H. retardatus salamander larvae, partly due to their lower toxin resistance and their higher toad consumption rate. Most importantly, omnivorous foraging traits (i.e., food sharing and carcass consumption) demonstrated by R. pirica frog tadpoles, but not by the strictly carnivorous H. retardatus salamander larvae, rendered the former particularly vulnerable to the toxic B. japonicus formosus toad hatchlings. This is the first study suggesting that species‐specific foraging traits such as food‐sharing and carcass consumption can determine predator vulnerability to toxic alien prey. This study emphasises the importance of investigating a wider array of predator traits as factors that determine their vulnerability to toxic alien prey. By doing so, we can understand what types of native species are strongly impacted by toxic alien prey and predict the ecological consequences of toxic species invasion.

An interaction-driven cannibalistic reaction norm.

sCannibalism is induced in larval‐stage populations of the Hokkaido salamander, Hynobius retardatus, under the control of a cannibalism reaction norm. Here, I examined phenotypic expression under the cannibalism reaction norm, and how the induction of a cannibalistic morph under the norm leads to populational morphological diversification. I conducted a set of experiments in which density was manipulated to be either low or high. In the high‐density treatment, the populations become dimorphic with some individuals developing into the cannibal morph type. I performed an exploratory analysis based on geometric morphometrics and showed that shape characteristics differed between not only cannibal and noncannibal morph types in the high‐density treatment but also between those morph types and the solitary morph type in the low‐density treatment. Size and shape of cannibal and noncannibal individuals were found to be located at either end of a continuum of expression following a unique size–shape integration rule that was different from the rule governing the size and shape variations of the solitary morph type. This result implies that the high‐density‐driven inducible morphology of an individual is governed by a common integration rule during the development of dimorphism under the control of the cannibalism reaction norm. Phenotypic expression under the cannibalism reaction norm is driven not only by population density but also by social interactions among the members of a population: variation in the populational expression of dimorphism is associated with contingent social interaction events among population members. The induced cannibalistic morph thus reflects not only by contest‐type exploitative competition but also interference competition.

Giant cannibals drive selection for inducible defence in heterospecific prey

Predation can strongly influence the evolution of prey species by selecting for defensive phenotypes that reduce the risk of predation. Although cannibalism is frequent in predators and is known to strongly influence predator–prey dynamics, it is largely neglected when studying the evolution of phenotypic defences in prey. Changes in cannibalism can alter the abundance, size-structure, and even phenotypes within predator populations, and thereby change the numerical and per-capita effects driving the selection pressure on defensive phenotypes in prey. In the present study, we experimentally manipulated the presence and absence of cannibalism in predatory salamander larvae Hynobius retardatus (Dunn) and examined how this affects the expression of and selection pressure on inducible defensive phenotype (bulgy phenotype) in its prey, Rana pirica (Matsui) tadpoles. In the absence of cannibalism, salamanders were generally not sufficiently large to consume tadpoles, tadpoles did not express the defensive phenotype, and all phenotypes had equally high survival. By contrast, cannibalism of salamanders accelerated growth rates, leading to the development of ‘giant’ cannibalistic morphs that were able to consume tadpoles. Importantly, only in the presence of these giant cannibals did tadpoles express a defensive ‘bulgy’ phenotype that had significantly higher survival rates than phenotypes without this defence, indicating that cannibalism altered the selective regime for defensive traits in prey. The results of the present study demonstrate that cannibalism can drive trait evolution in predator–prey systems and suggest that we cannot predict how changes in numerical effects in predator populations affect selection regimes on prey traits without accounting for concurrent changes in predator per-capita effects.

Contacts with large, active individuals intensify the predation risk of small conspecifics.

Size variation within a population can influence the structure of ecosystem interactions, because ecological performance differs between individuals of different sizes. Although the impact of size variation in a predator species on the structure of interactions is well understood, our knowledge about how size variation in a prey species might modify the interactions between predators and prey is very limited. Here, by examining the interactions between predatory Hynobius retardatus salamander larvae and their prey, Rana pirica frog tadpoles, we investigated how large prey individuals affect the predation mortality of small prey conspecifics. First, in an experiment conducted in a field pond in which we manipulated the presence of salamanders and large tadpoles (i.e., large enough to protect them against salamander predation) with small tadpoles, we showed that in the presence of large tadpoles the mortality of small tadpoles from salamander predation was increased. On the basis of our observations of the activity of individuals, we hypothesized that active large tadpoles caused physical disturbances, which in turn caused the small tadpoles to move, and thus increased their encounter frequency with the predatory salamanders. To test this hypothesis, we conducted a laboratory experiment in small tanks with three players (i.e., one salamander as predator, one small tadpole as focal prey, and either a small or a large tadpole as the prospective movement inducer). In each tank, we manipulated the presence or absence of a movement inducer, and, when present, its size (large or small) and access (caged or uncaged) to the focal prey. In the presence of a large, uncaged movement inducer, the focal prey was more active and suffered from higher predation mortality compared with the other treatments, because the large movement inducer (unlike a small movement inducer) moved actively and, when uncaged, could stimulate movement of the focal prey through direct contact. The results indicated that high activity of large prey individuals and the resulting behavioral interactions with small conspecifics via direct contact indirectly increased the mortality of the small prey.

Retardation of larval development in the salamander Hynobius retardatus in a permanent pond with abundant spring water

Plasticity in the larval life history of the Ezo salamander Hynobius retardatus has been reported. In the present study, we monitored larvae of this salamander species in a fragmented forest in the environs of Sapporo, Japan. Overwintering larvae were detected in one of the two ponds examined, i.e., in the pond that was permanent, with water supplied from several spring-fed points. Four seasonal transition phases in water temperature were observed because of the abundant spring-fed groundwater supply and canopy cover from May to October. These phases included a warming period, a constant-high period, a cooling period, and a constant-low period. During the constant-low period, premetamorphic larvae that had already emerged during the cooling period were continuously detected; however, the composition of the developmental stages remained unchanged, with larval growth progressing slowly. There is apparently a critical temperature that represents the threshold for metamorphosis initiation. The critical temperature is expected to be slightly higher than the groundwater temperature at the spring-fed points. Emigration of overwintered larvae resumed during the warming period and continued during the constant-high period in the year following hatching. In the nearby temporary pond, the one-year-old cohort completed metamorphosis during the summer of the hatch year.

An aquatic vertebrate can use amino acids from environmental water.

Conventional food-web theory assumes that nutrients from dissolved organic matter are transferred to aquatic vertebrates via long nutrient pathways involving multiple eukaryotic species as intermediary nutrient transporters. Here, using larvae of the salamander Hynobius retardatus as a model system, we provide experimental evidence of a shortcut nutrient pathway by showing that H. retardatus larvae can use dissolved amino acids for their growth without eukaryotic mediation. First, to explore which amino acids can promote larval growth, we kept individual salamander larvae in one of eight different high-concentration amino acid solutions, or in control water from which all other eukaryotic organisms had been removed. We thus identified five amino acids (lysine, threonine, serine, phenylalanine, and tyrosine) as having the potential to promote larval growth. Next, using 15N-labelled amino acid solutions, we demonstrated that nitrogen from dissolved amino acids was found in larval tissues. These results suggest that salamander larvae can take up dissolved amino acids from environmental water to use as an energy source or a growth-promoting factor. Thus, aquatic vertebrates as well as aquatic invertebrates may be able to use dissolved organic matter as a nutrient source.

Population Structure and Evolution after Speciation of the Hokkaido Salamander (Hynobius retardatus).

The Hokkaido salamander (Hynobius retardatus) is endemic to Hokkaido Island, Japan, and shows intriguing flexible phenotypic plasticity and regional morphological diversity. However, to date, allozymes and partial mitochondria DNA sequences have provided only an outline of its demographic histories and the pattern of its genetic diversification. To understand the finer details of the population structure of this species and its evolution since speciation, we genotyped five regional populations by using 12 recently developed microsatellite polymorphic markers. We found a clear population structure with low gene flow among the five populations, but a close genetic relationship between the Teshio and Kitami populations. Our demographic analysis suggested that Teshio and Erimo had the largest effective population sizes among the five populations. These findings regarding the population structure and demography of H. retardatus improve our understanding of the faunal phylogeography on Hokkaido Island and also provide fundamental genetic information that will be useful for future studies.

Pseudoacanthocephalus toshimai sp. nov. (Palaeacanthocephala: Echinorhynchidae), a common acanthocephalan of anuran and urodelan amphibians in Hokkaido, Japan, with a finding of its intermediate host

The Ezo brown frog (Rana pirica) and the Ezo salamander (Hynobius retardatus) are endemic species of Hokkaido, the northernmost island of Japan. Intestinal adult acanthocephalans are common in these amphibians. A molecular identification based on nuclear and mitochondrial DNA markers demonstrated that the parasites from the anuran and the urodelan are the same species. In the neighboring Honshu island, another acanthocephalan from ranid frogs (e.g. Rana japonica and Rana ornativentris) has been identified as Acanthocephalus lucidus. The counterpart species from the amphibians of Hokkaido was morphologically indistinguishable from A. lucidus. However, clear genetic distinctiveness between the two allopatric populations (separated by islands) indicated the entity of a cryptic species. A phylogenetic tree inferred from sequences of 28S ribosomal DNA showed that the acanthocephalans from Honshu and Hokkaido belong to the genus Pseudoacanthocephalus. Therefore, Pseudoacanthocephalus toshimai sp. nov. is proposed for the cryptic species in Hokkaido, together with the transfer of A. lucidus in Honshu to Pseudoacanthocephalus lucidus comb. nov. The present field survey further demonstrated Ligidium japonicum, an isopod crustacean living in the litter layer of forests, to be an intermediate host of the new species.

Development and Characterization of 12 Microsatellite Markers for the Hokkaido Salamander (Hynobius retardatus)

Abstract: The Hokkaido salamander, Hynobius retardatus (Caudata: Hynobiidae), is a common salamander species distributed throughout Hokkaido Island, Japan. Because of its regional morphological diversity and flexible phenotypic plasticity, its demographic history and the causes of its distribution patterns are of intrinsic interest to evolutionary biologists. To address these issues, a microsatellite marker is an ideal genetic marker for inferring gene flows and hybridizations between populations on a fine scale. We therefore utilized an Ion PGM™ sequencing system to screen the H. retardatus genome for microsatellite markers. As the result, we identified 12 polymorphic markers. We then tested the usefulness of these markers in H. retardatus by genotyping 20 individuals from each of two regional populations (the Erimo and Nopporo populations). The total number of alleles ranged from 3 to 8, and the expected heterozygosities per locus ranged from 0.049 to 0.803. We expect the developed markers to be useful ...

Predator cannibalism can intensify negative impacts on heterospecific prey.

Although natural populations consist of individuals with different traits, and the degree of phenotypic variation varies among populations, the impact of phenotypic variation on ecological interactions has received little attention, because traditional approaches to community ecology assume homogeneity of individuals within a population. Stage structure, which is a common way of generating size and developmental variation within predator populations, can drive cannibalistic interactions, which can affect the strength of predatory effects on the predator's heterospecific prey. Studies have shown that predator cannibalism weakens predatory effects on heterospecific prey by reducing the size of the predator population and by inducing less feeding activity of noncannibal predators. We predict, however, that predator cannibalism, by promoting rapid growth of the cannibals, can also intensify predation pressure on heterospecific prey, because large predators have large resource requirements and may utilize a wider variety of prey species. To test this hypothesis, we conducted an experiment in which we created carnivorous salamander (Hynobius retardatus) populations with different stage structures by manipulating the salamander's hatch timing (i.e., populations with large or small variation in the timing of hatching), and explored the resultant impacts on the abundance, behavior, morphology, and life history of the salamander's large heterospecific prey, Rana pirica frog tadpoles. Cannibalism was rare in salamander populations having small hatch-timing variation, but was frequent in those having large hatch-timing variation. Thus, giant salamander cannibals occurred only in the latter. We clearly showed that salamander giants exerted strong predation pressure on frog tadpoles, which induced large behavioral and morphological defenses in the tadpoles and caused them to metamorphose late at large size. Hence, predator cannibalism arising from large variation in the timing of hatching can strengthen predatory effects on heterospecific prey and can have impacts on various, traits of both predator and prey. Because animals commonly broaden their diet as they grow, such negative impacts of predator cannibalism on the heterospecific prey may be common in interactions between predators and prey species of similar size.

Antagonistic indirect interactions between large and small conspecific prey via a heterospecific predator

Intrapopulation size variation strongly influences ecological interactions because individuals belonging to different size groups have distinct functions. Most demonstrations of the impacts of size variation in trophic systems have focused on size variation in predator species, and the consequences of size variation in prey species are less well understood. We investigated how prey size structure shapes intra- and interspecific interactions in experiments with a gape-limited predator (larvae of the salamander Hynobius retardatus) and its heterospecific prey (frog tadpoles, Rana pirica). We found that large and small tadpole size groups each increased mortality in the other group by intensifying salamander predation; this type of indirect interactions is called apparent competition. The antagonistic impacts on the prey size groups were caused by different size-specific mechanisms. By consuming small tadpoles, the salamanders grew large enough to consume large tadpoles. The activity of large tadpoles, by increasing the activity of the small tadpoles, may increase the number of encounters with the predator and thus small tadpole mortality. These results suggest that the magnitude of a predator's ecological role, such as whether a top–down trophic cascade is initiated, depends on size variation in its heterospecific prey.

Transcriptome analysis of predator- and prey-induced phenotypic plasticity in the Hokkaido salamander (Hynobius retardatus).

Predator- and prey-induced phenotypic plasticity is widely observed among amphibian species. Although ecological factors inducing diverse phenotypic responses have been extensively characterized, we know little about the molecular bases of variation in phenotypic plasticity. Larvae of the Hokkaido salamander, Hynobius retardatus, exhibit two distinct morphs: the presence of their prey, Rana pirica tadpoles, induces a broad-headed attack morph, and the presence of predatory dragonfly nymphs (Aeshna nigroflava) induces a defence morph with enlarged external gills and a high tail. To compare the genes involved in predator- and prey-induced phenotypic plasticity, we carried out a de novo transcriptome analysis of Hokkaido salamander larvae exposed to either prey or predator individuals. First, we found that the number of genes involved in the expression of the defence morph was approximately five times the number involved in the expression of the attack morph. This result is consistent with the fact that the predator-induced plasticity involves more drastic morphological changes than the prey-induced plasticity. Second, we found that particular sets of genes were upregulated during the induction of both the attack and defence morphs, but others were specific to the expression of one or the other morph. Because both shared and unique molecular mechanisms were used in the expression of each morph, the evolution of a new plastic phenotype might involve both the co-option of pre-existing molecular mechanisms and the acquisition of novel regulatory mechanisms.