COMPETITION AMONG TADPOLES OF COEXISTING HEMICLONES OF HYBRIDOGENETIC RANA ESCULENTA: SUPPORT FOR THE FROZEN NICHE VARIATION MODEL

Vertebrate animals reproducing without genetic recombination typically are hybrids, which have large ranges, are locally abundant, and live in disturbed or harsh habitats. This holds for the hemiclonal hybridogenetic frog Rana esculenta: it is widespread in Europe and commonly is found in disturbed habitats such as gravel pits. We hypothesize that its widespread occurrence may either be the result of natural selection for a single hemiclone acting as a broadly adapted “general-purpose” genotype, or of interclonal selection, which maintains multiple hemiclones that each are relatively narrowly adapted and perform differently across environments, that is, the Frozen Niche Variation model. We tested these competing hypotheses using 1000-L outdoor artificial ponds to rear tadpoles of the parental species (Rana lessonae [LL] and Rana ridibunda [RR]) alone, and each of three hemiclones of Rana esculenta (GUT1, GUT2, GUT3) alone, and in mixed hemiclonal populations from hatching to metamorphosis. Tadpoles of three coexisting hemiclones from a single natural population (near Gütighausen, Switzerland) were reared in both two- and three-way mixtures in equal total numbers at high and low density. For each species and hemiclone, the proportion of tadpoles metamorphosing decreased as the density of tadpoles increased, with the three hemiclones spanning the range of values exhibited by the two parental species. LL and GUT1 tadpoles produced the highest proportion of metamorphs, whereas tadpoles of RR produced the fewest metamorphs at both densities. GUT1 tadpoles also produced the largest metamorphs at low density, GUT2 and GUT3 tadpoles produced smaller metamorphs than did GUT1 tadpoles at the low density, but the three hemiclones did not differ from each other at high density. The parental species (LL and RR) were intermediate in metamorphic size to the hemiclones at low density, but all genotypes converged on a similar size at high density. Length of the larval period also was affected by density, but its effect was dependent on genotype. GUT1 tadpoles had the shortest larval period at the low density, but larval period was longer and not different between GUT1, GUT3, and LL at high density. RR tadpoles had the longest larval period at both densities. The most dramatic results were that three genotypes (GUT1, GUT2, and RR) maintained rank order and increased days to metamorphosis from low to high density, whereas two genotypes (GUT3 and LL) changed rank order and decreased days to metamorphosis from low to high density. Mixtures of hemiclones in two- and three-way combinations facilitated the proportion of tadpoles metamorphosing for GUT1 and GUT2 at both densities, but only at the low density for GUT3 tadpoles. Results from this experiment are incompatible with the General-Purpose Genotype model as a global explanation of hybrid abundance in these frogs. Alternatively, the Frozen Niche Variation prediction of general performance superiority of clonal mixtures relative to single clone populations is strongly supported. The data confirm that fitness advantages of hemiclones change, depending on the environment, such that in temporally and spatially heterogeneous habitats like ponds, frequency-dependent selection among hemiclones may promote coexistence in hemiclonal assemblages. Yet, differential dispersal or colonization ability and historical factors affecting hemiclone distribution may also be important in shaping patterns of clonal coexistence.     

ASYMMETRIC COMPETITION IN MIXED POPULATIONS OF TADPOLES OF THE HYBRIDOGENETIC: RANA ESCULENTA COMPLEX

Hybridogenetic Rana esculenta tadpoles display tolerance to extreme environmental conditions and fit criteria of the “general-purpose” genotype. A trade-off between generality and competitive ability is assumed to occur in asexual species, but the evidence remains unclear. The purpose of my experiment was to test the competitive ability of hemiclonal hybrid Rana esculenta tadpoles relative to the parental species Rana lessonae. Mixed and single genotype populations of R. esculenta and R. lessonae tadpoles were reared at three densities in artificial ponds. Survival of R. esculenta was higher than for R. lessonae tadpoles, but did not differ among densities. Body size at metamorphosis was the same between genotypes, but decreased with increasing density. Larval period was not affected by density, but R. esculenta tended to metamorphose earlier than R. lessonae. Percentage of individuals metamorphosing was higher for R. esculenta at both medium and high densities, but the same as R. lessonae at the low density. The difference in survival, body size, and larval period between tadpoles reared in single and mixed genotype populations was unaffected by genotype or density. The difference in the percentage of metamorphs, however, was strongly affected. The percentage of hybrids metamorphosing was 9% above the responses of single genotype populations at the highest density. Conversely, the percentage of R. lessonae metamorphosing was 12% below the responses of single genotype populations at the same density. Hybrid success in this experiment further supports the criterion of a “general-purpose” genotype without assumptions of reduced competitive ability.     

Effects of density and predation on Scaphiopus couchi tadpoles in desert ponds

Summary The effects of density on growth and development of Scaphiopus couchi tadpoles in desert ponds were investigated, and sources of mortality over a three-year period were documented. In 16 of the 82 ponds monitored, predation was the principal cause of death, demonstrating that tadpoles in desert ponds may be exposed to high levels of predation, although the overall importance of predation is less here than in more mesic areas. Desiccation was the primary cause of mortality in 49 ponds. Growth and development were extremely slow in most high density ponds and as a result most tadpoles were unable to metamorphose before ponds dried. Only 8 ponds produced metamorphs, and mortality was high even in these. Food-supplementation resulted in some metamorphosis in high density ponds, although the effect was diminished by the extreme crowding in most ponds. In low density ponds, S. couchi tadpoles can develop very quickly and metamorphose. High mortality due to desiccation is largely a consequence of high density: tadpoles rarely completed development in high density ponds, regardless of pond duration.     

Rana temporaria metamorph production and population dynamics in the field – Effects of tadpole density,predation and pond drying

This study investigates factors of importance for tadpoles survival and metamorph production in the common frog Rana temporaria. It also assess the importance of this for the population dynamics of the species. Eighteen ponds were studied for up to 8 years. Data collected each year included: number of spawn clumps deposited, tadpole number and metamorph number. The permanency of the ponds was also recorded each year. Measures were taken of predator density. There was no suggestion of density dependence in the survival of tadpoles. In contrast, the number of spawn clumps deposited per pond area was highest for ponds with high survival. Density of predators (sticklebacks, newts and invertebrates) was negatively correlated to tadpole and metamorph survival. This was true both within (among years) and among ponds. Several of the study ponds dried completely before metamorphosis in some years. However, those ponds also were those with the smallest number of predators and in years with successful metamorphosis, these ponds produced more metamorphs than more permanent ponds. An analysis of the year to year dynamics showed that population size (number of deposited spawn clumps) was correlated to that in the previous year, suggesting a fairly high adult survival, but also on the number of metamorphs emerging two or three years before (corresponding to the age of sexual maturity of the species). It is concluded that the aquatic stage is not strongly limiting in these ponds but conservation efforts should be focused on the terrestrial habitat. Also, the study stresses the value of temporary ponds, despite the fact that recruitment often fails totally in these.     

Tadpole competition in a desert anuran community

Summary Five interspecific competition experiments performed in artificial ponds under temporary pond-like conditions established that temporary and permanent pond tadpoles interact. Temporary pond species typically lowered survivorship and reduced progress towards metamorphosis of permanent pond tadpoles. These results suggest that competitive interactions may partially determine the local distribution of tadpoles in the Chihuahuan Desert by preventing some permanent pond anurans from using temporary ponds. Predation by Scaphiopus multiplicatus tadpoles (a temporary pond species) may also prevent permanent pond tadpoles from using temporary ponds.     

POLYPHENISM IN SPADEFOOT TOAD TADPOLES AS A LOCALLY ADJUSTED EVOLUTIONARILY STABLE STRATEGY

I examined the evolutionary factors maintaining two environmentally induced morphs in ponds of variable duration. Larvae of New Mexico spadefoot toads (Scaphiopus multiplicatus) often occur in the same pond as a large, rapidly developing carnivorous morph and as a smaller, more slowly developing omnivorous morph. Previous studies revealed that carnivores can be induced by feeding tadpoles live fairy shrimp and that morph determination is reversible. Field and laboratory experiments indicated that the ability of an individual to become a carnivore or an omnivore is maintained evolutionarily as a response to variability in pond longevity and food abundance. Carnivores survived better in highly ephemeral artificial ponds, because they developed faster. Omnivores survived better in longer-duration artificial ponds, because their larger fat reserves enhanced postmetamorphic survival. The two morphs also occupy different trophic niches. Experimental manipulations of morph frequency in ponds of intermediate duration revealed that increased competition for food among individuals of the more common morph made the rarer form more successful. Morph frequency within each pond was stabilized at an equilibrium by frequency-dependent morph reversal, which reflected frequency-dependent natural selection on size at metamorphosis: larger metamorphs had higher survival, and individuals reared at a frequency above the pond's equilibrium frequency were smaller at metamorphosis than were individuals of that morph reared at a frequency below the pond's equilibrium. Because neighboring ponds often differed in pond longevity and food abundance, each pond possessed a unique equilibrium morph frequency. This implies that morph determination in Scaphiopus is a locally adjusted evolutionarily stable strategy (ESS).     

Canopy Closure,Competition, and the Endangered Dusky Gopher Frog

Abstract: A major challenge facing wildlife biologists is understanding why some species go extinct while others persist in the same habitat. To address this question, we investigated whether tree canopy closure over ponds affects growth and survival of rare and common tadpoles within ponds and mediates competitive interactions among species. We conducted 2 experiments to test whether canopy closure and competition may have contributed to the decline of the endangered dusky gopher frog (Rana sevosa), but allowed the persistence of the southern leopard frog (R. sphenocephala). We explored the response of both species to canopy closure in single-species and mixed- (1:1) species treatments of identical total tadpole density. An experiment using aquatic enclosures in temporary ponds showed that canopy closure reduced tadpole growth approximately 20% for both species. Survival of dusky gopher frog tadpoles was higher in mixed-species enclosures than in single-species enclosures. In a complementary experiment using artificial ponds, dusky gopher frogs had lower survival to metamorphosis, reduced size at metamorphosis, and produced a lower total biomass of metamorphosed juveniles in shaded ponds. Southern leopard frogs exhibited reduced body size at metamorphosis only when shaded. These studies suggest that pond canopy closure, not larval competition, may be contributing to the decline of the dusky gopher frog. The different responses to canopy closure suggest a potential mechanism for the loss of dusky gopher frogs and the persistence of southern leopard frogs. Removal of trees from historically open-canopy ponds may help facilitate the recovery of dusky gopher frogs and benefit similar species.     

Differential performance among LDH-B genotypes in Rana lessonae tadpoles

The European pool frog, Rana lessonae, is widely polymorphic for two common alleles (b,e) at the lactate dehydrogenase-B (LDH-B) locus. We compared fitness-related larval life-history traits among LDH-B genotypes, which originated from segregation in heterozygous parents, in an artificial pond experiment where tadpoles of R. lessonae from a Swiss population were raised together with tadpoles of the hemiclonal hybrid R. esculenta at two densities. In R. lessonae, LDH-B e/e homozygotes at each density had a higher proportion of metamorphs among survivors, reached metamorphosis earlier, and were heavier at metamorphosis than b/b homozygotes; b/e heterozygotes had intermediate values. That e/e individuals were superior to b/b in both time to and mass at metamorphosis is surprising because these two life-history traits are thought to reflect a performance trade-off; e/e genotypes apparently compensated for shorter time to metamorphosis by a higher growth rate. The two alleles showed the same performance ranking when combined in hybrids with a R. ridibunda allele: When R. esculenta from Swiss populations reared in the same ponds had received the e allele rather than the b allele from their R. lessonae parent, they reached metamorphosis earlier, but did not differ in mass at metamorphosis. The degree of linkage disequilibrium in the source population of the eight R. lessonae used as parents of the R. lessonae tadpoles is unknown, so we cannot exclude the possibility that the performance differences are caused by some anonymous tightly linked gene, rather than the LDH-B locus, that constitutes the genomically localized target of natural selection. A causal involvement of LDH-B is plausible, nevertheless, because this enzyme takes part in the central energy-metabolizing processes and has been reported to underlie fitness differences in other animals; also, differential performance of LDH-B genotypes has been observed in R. lessonae larvae from another population. The present results suggest strong directional selection for allele e; the sum of available data, including an independent laboratory experiment, suggests that partial environment-dependent overdominance combined with balancing selection favoring e/e homozygotes under some and b/b homozygotes under other conditions may be partially responsible for the broad maintenance of the LDH-B polymorphism in R. lessonae.     

Experimental analysis of the evolutionary potential of hybridization in leopard frogs (Anura: Ranidae)

Artificial crossing using Rana blairi and R. sphenocephala frogs produced conspecific, interspecific and F₁ backcross hybrid genotypes. Although hybrid males used in the crosses were sterile, crosses using hybrid females produced viable larvae. The larval performance of resultant parental and hybrid genotypes was measured in experimental ponds at two densities. Density significantly affected survival, body mass at metamorphosis, larval period length and metamorphosis for all genotypes. Survival was the same among genotypes, but decreased with increasing density. Body mass at metamorphosis was the same among genotypes, but decreased with increasing density. Larval period increased with increasing density. Among genotypes, larvae from the conspecific R. sphenocephala cross had the shortest larval period while larvae from the conspecific R. blairi cross had the longest larval period. All hybrid genotypes had larval periods longer than R. sphenocephala, but shorter than R. blairi. The percentage of individuals metamorphosing was highest for R. sphenocephala ponds and lowest for R. blairi ponds across densities. Ponds with hybrid larvae produced a greater proportion of metamorphs than those with R. blairi larvae, but a smaller proportion than R. sphenocephala ponds. Equivalent or increased relative larval performance of hybrid genotypes under the conditions of our experiment suggests that hybrid genotypes may possess similar or higher fitnesses relative to their progenitors in some environments. Reduced fertility of adult hybrid males is a powerful selective force against natural hybridization. Nevertheless, because of the successful reproduction by female hybrids, natural hybridization has the potential to serve as a mechanism for the introgression of novel genetic variation that can benefit both R. blairi and R. sphenocephala in fluctuating and unpredictable larval environments. Experimental determination of the fitness of parental and hybrid genotypes is crucial for a comprehensive understanding of the effects of hybridization on organismal evolution.     

FIXATION OF DELETERIOUS MUTATIONS IN CLONAL LINEAGES: EVIDENCE FROM HYBRIDOGENETIC FROGS

Abstract The hemiclonal waterfrog Rana esculenta (RL genotype), a bisexual hybrid between R. ridibunda (RR) and R. lessonae (LL), eliminates the L genome from its germline and clonally transmits the R genome (hybridogenesis). Matings between hybrids produce R. ridibunda offspring, but they generally die at an early larval stage. Mortality may be due to fixed recessive deleterious mutations in the clonally inherited R genomes that were either acquired through the advance of Muller's ratchet or else frozen in these genomes at hemiclone formation. From this hypothesis results a straightforward prediction: Matings between different hemiclones, that is, between R. esculenta possessing different R genomes of independent origin, should produce viable R. ridibunda offspring because it is unlikely that different clonal lineages have become fixed for the same mutations. I tested this prediction by comparing survival and larval performance of tadpoles from within‐ and between‐population crossings using R. esculenta from Seseglio (Se) in southern, Alpnach (Al) in central, and Elliker Auen (El) in northern Switzerland, respectively. Se is isolated from the other populations by the Alps. Enzyme electrophoresis revealed that parents from Se belonged to a single hemiclone that was different from all hemiclones found north of the Alps. Parents from Al also belonged to one hemiclone, but parents from El belonged to three hemiclones, one of which was indistinguishable from the one in Al. Rana esculenta from Se produced inviable tadpoles when crossed with other hybrids of their own population, but when crossed with R. esculenta from Al and El, tadpoles successfully completed metamorphosis, supporting the hypothesis I tested. Within‐population crosses from Al were also inviable, but some within‐population crosses from El, where three hemiclones were present, produced viable offspring. Only part of the crosses between Al and El were viable, but there was no consistent relationship between hemiclone combination and tadpole survival. When backcrossed with the parental species R. ridibunda, hybrids from all source populations produced viable offspring. Performance of these tadpoles with a sexual and a clonal genome was comparable to that of normal, sexually produced R. ridibunda tadpoles. Thus, in the heterozygous state, the deleterious mutations on the clonal R genomes did not appear to reduce tadpole fitness.     

The formation of new characteristics in life cycle of the marsh frog (Rana ridibunda) in thermal pond conditions

Using mark and recapture approach, the long-term population dynamics in the marsh frog (Rana ridibunda) was studied. Group-marking of metamorphs was conducted in a small thermal pond serving as a sedimentation basin for discharged waters from Nizhny Tagil metallurgic works. Depending on the time of metamorphosis, three groups of individuals could be singled out, namely: early ones (group I), middle ones (group II), and late ones that overwinter as tadpoles and complete metamorphosis in May of the next year (group III). Upon metamorphosis completion, individuals of group I were found to be significantly larger than those of group II, and individuals of both these groups to be significantly smaller than those of group III. After first wintering, immature individuals from group I were significantly larger than either individuals from group II or metamorphs from group III, though a growth rate of the latter was significantly higher than in groups I and II. These discrepancies were observed both between immature and adult individuals. Over the period from metamorphosis completion to the first wintering ending, survivorship in group I was significantly higher and did not differ between groups II and III. In adult frogs, maximum survivorship was registered in group III and minimum one in group II; the detected differences recurred in each age class till the fourth wintering. However, in age classes that overwintered 4 and 5 times, maximum survivorship was observed in group II, which can be treated as a compensation for rather low survivorship of this group at younger ages. All the events of tadpoles of this species overwintering (except in other thermal water bodies) that are described in literature, correspond to rare deviations from normal ontogenesis. Therefore, the revealed formation of a numerous group of overwintering tadpoles in successive generations should be considered as a new adaptation which sense is a decrease of competition between tadpole groups when using the highly productive resources of the thermal pond practically year-round. The advantage in body size and growth rate of not only tadpoles but also of metamorphs, immature and adult individuals of group III indicates that after metamorphosis the strategy of this group still remains successful. The reason for that is unusually large body size of metamorphs which provides higher postmetamorphic survivorship and greater female fecundity.     

Growth and development of larval green frogs (Rana clamitans) exposed to multiple doses of an insecticide

Our objective was to determine how green frogs (Rana clamitans) are affected by multiple exposures to a sublethal level of the carbamate insecticide, carbaryl, in outdoor ponds. Tadpoles were added to 1,000-l ponds at a low or high density which were exposed to carbaryl 0, 1, 2, or 3 times. Length of the larval period, mass, developmental stage, tadpole survival, and proportion metamorphosed were used to determine treatment effects. The frequency of dosing affected the proportion of green frogs that reached metamorphosis and the developmental stage of tadpoles. Generally, exposure to carbaryl increased rates of metamorphosis and development. The effect of the frequency of carbaryl exposure on development varied with the density treatment; the majority of metamorphs and the most developed tadpoles came from high-density ponds exposed to carbaryl 3 times. This interaction suggests that exposure to carbaryl later in the larval period stimulated metamorphosis, directly or indirectly, under high-density conditions. Our study indicates that exposure to a contaminant can lead to early initiation of metamorphosis and that natural biotic factors can mediate the effects of a contaminant in the environment.     

Primary and secondary phenology. Does it pay a frog to spawn early?

This study examines the consequences of variation in the laying and hatching date for the time of metamorphosis in the common frog Rana temporaria . Field data are presented showing that eggs laid early tend to take longer to develop. Thus, the time advantage for early eggs is reduced at the time of hatching. There was an among-year variation in this phenomenon; it was not manifest in a phenologically late year. Also, field data revealed that mortality due to pond freezing is a real risk for early laid eggs. Finally, two experiments in tanks analyse the effects of hatching date variation for the time of metamorphosis. (1) When hatching was experimentally delayed by 7 or 11 days, this resulted in later metamorphosis, however, by only 2 and 5 days, respectively. (2a) When tadpoles from the same pond that naturally hatched at different times were compared, it was found that a hatching time difference of 6 days resulted in later metamorphosis by 2 days only. (2b) A comparison of tadpoles from two different ponds that hatched 11 days apart also resulted in only 2 days' difference in metamorphosis. In this case, the later but faster developing tadpoles metamorphosed at a smaller size. I suggest that eggs from these two ponds differed genetically in the growth and development strategy. Despite the obvious risks, and the moderate gain in terms of early metamorphosis, frogs breed dangerously early in spring. Possible reasons for this are discussed. These include external selective forces that promote early metamorphosis (also at a high cost), within-pond competition among tadpoles with an advantage for early and large tadpoles and finally factors relating to mate choice at the breeding site.     

DELAYED COSTS OF AN INDUCED DEFENSE IN TADPOLES? MORPHOLOGY,HOPPING, AND DEVELOPMENT RATE AT METAMORPHOSIS

Models for the evolution of plasticity predict that individuals having phenotypes induced by exposure to enemies should experience relatively low fitness when enemies are absent. However, costs of induced phenotypes have been difficult to find in both plants and animals, perhaps because costs are expressed at later stages in the life cycle. We searched for delayed costs of an induced defense in larvae of the water frog Rana ridibunda, which exhibits strong phenotypic responses to predators. Tadpoles grew to metamorphosis in outdoor artificial ponds, in either the presence or absence of Aeshna dragonfly larvae confined within cages. We collected metamorphs at forelimb emergence, estimated their development rate until tail resorption was complete, and measured their body and leg shape and hopping performance. Development rate through metamorphosis reflects the duration of a transitional period during which metamorphs are especially vulnerable to predators, and hopping performance may reflect ability to escape predators. Froglets from the dragonfly treatment lost mass through metamorphosis significantly faster than those from predator-free ponds, but they resorbed their tails at about the same rate, despite the fact that their tails were relatively large to begin with. Froglets developing from predator-induced tadpoles had shorter, more muscular legs, and hopped 5% longer distances (difference not significant). Therefore, producing an induced defense against insect predators during the tadpole stage did not exact a cost during or immediately after metamorphosis; if anything, tadpoles with the predator-induced phenotype gave rise to more vigorous froglets. These results focus attention on other costs of the induced phenotype, as well as alternative explanations for plasticity that do not rely on direct fitness trade-offs.     

Competing tadpoles: Australian native frogs affect invasive cane toads (Rhinella marina) in natural waterbodies

The cane toad (Rhinella marina) is one of the most successful invasive species worldwide, and has caused significant negative impacts on Australian fauna. Experimental work in the laboratory and in mesocosms has shown that tadpoles of native frogs can affect survival, size at metamorphosis and duration of larval period of cane toad tadpoles. To test if these effects occur in nature, we conducted a field experiment using two temporary ponds where we set up enclosures with tadpoles of native green tree frogs (Litoria caerulea) and cane toads in treatments with a range of densities and combinations. The presence of green tree frog tadpoles significantly decreased the growth rate of toad tadpoles and increased the duration of their larval period in both ponds; in one pond, frog tadpoles also significantly reduced the body length and mass of metamorph toads. Toad tadpoles did not have any significant negative effects on green tree frog tadpoles, but there was strong intraspecific competition within the latter species: increased frog tadpole density resulted in increased larval period and reduced survival, growth rate and size at metamorphosis for frogs at one or both ponds. Our results are encouraging for the possibility of using native frogs as one component of an integrated approach to the biological control of cane toads.     

Influence of density and predators on metamorphic size in Rhinella schneideri tadpoles raised in mesocosm conditions

Amphibians exhibit extreme plasticity in the timing of metamorphosis, and several species respond to water availability, accelerating metamorphosis when their ponds dry. We analyzed the plasticity of the developmental response to water volume in Rhinella schneideri tadpoles. We raised tadpoles in mesocosm. Covariation between body size at metamorphosis and timing of development was positive. Nevertheless, the first approximately 53% of the metamorphoses finishing the cycle required between 34 and 56 days, and the covariation between body size at metamorphosis and timing of development was negative. For these tadpoles, the larval density and the presence of predators did not significantly affect their mass to metamorphosis. Nevertheless, predators affected time to metamorphosis. For the remainder of the tadpoles that reached metamorphosis at > 56 days, the relationship between body size at metamorphosis and timing of development was positive. For these tadpoles, larval density was important for mass at metamorphosis and presence of predators was also important for time to metamorphosis. Two dominant features were observed: (i) approximately 53% of metamorphs had morphological features similar to individuals developing in desiccating ponds, and (ii) the other individuals had morphological characteristics comparable to metamorphs developing in an unchanging environment.     

Male vocalization and female choice in the hybridogenetic Rana lessonae/Rana esculenta complex

In many species, females can improve their fitness by preferring particular males over others. In Palaearctic water frogs of the Rana lessonae/R. esculenta complex the consequences of such mate choice are particularly pronounced. To produce viable offspring, the hybrid R. esculenta (genotype RL) must mate with the parental species R. lessonae (LL); but R. lessonae should avoid mating with R. esculenta, because the resulting hybrid offspring will eliminate the L genome from the germline (hybridogenesis). Hence, there exists a conflict between the sexual parasite (RL) and its sexual host (LL) over the best mating partner. Previous studies have shown a preference for LL males in LL and RL females; but they have also shown that females cannot usually realize their choice when in close proximity to males, because the males forcefully and indiscriminately amplex them. We tested whether females use male vocalizations as a long-distance signal to increase their chances of mating with the preferred LL males. We exposed female R. lessonae and R. esculenta to playbacks of single LL and RL mating calls (experiment 1) and to choruses with a 3:1 excess of LL and RL calls, respectively (experiment 2). In experiment 1, both female types were attracted more by the LL than by the RL calls. In experiment 2, no discrimination between LL- and RL-dominated choruses was observed. The results suggest that females do not use distant male vocalization to approach preferentially ponds or arenas within a pond that hold an excess of LL males. But once they have arrived in a chorus, mating calls from nearby males can direct them to the preferred LL mates. We discuss possible reasons for the failure to discriminate between choruses and the chances for successful choice between individuals within choruses. Copyright 2000 The Association for the Study of Animal Behaviour.     

ADAPTIVE PLASTICITY IN DEVELOPMENT OF SCAPHIOPUS COUCHII TADPOLES IN DESERT PONDS

Couch's spadefoot toads (Scaphiopus couchii) breed in ephemeral desert ponds that are highly variable in duration. Rapid development is expected to be advantageous in short-duration ponds, but slower development, allowing more time for growth, may be advantageous in ponds of longer duration. Previous experiments have revealed both genetic variation in development time and phenotypic plasticity in response to pond drying. In this paper, I examine the norms of reaction of five sibships of tadpoles to see whether there is genetic variation in the effect of pond duration, i.e., in phenotypic plasticity. Several important results emerged. 1) Differences among sibships in development time that were seen in the lab were also seen in the field. 2) There was no evidence for genetic variation in plasticity of development; all sibships exhibited faster development and decreased larval period in ponds of short duration. Plasticity in development appears to be adaptive, as size at metamorphosis was correlated with duration of larval period. The slowest developing sibship, however, suffered higher mortality compared to other sibships in short duration ponds. 3) Sibships did not differ in growth or size at metamorphosis in short-duration ponds, but the slowest developing sibship metamorphosed at the largest size in long duration ponds, resulting in a significant genotype x environment interaction for size at metamorphosis. Thus, although only one of the five sibships responded differently, there appears to be genetic variation for plasticity in growth, and a genetically determined trade-off between fitness in short-duration ponds (via rapid development) and fitness in long duration ponds (via large size at metamorphosis). This may explain the existence of both phenotypic plasticity and genetic variation in development. A single genotype, although capable of adaptive plasticity, is not sufficiently flexible to have equally high fitness in both long- and short-duration ponds.     

Heterozygous fitness effects of clonally transmitted genomes in waterfrogs

The European waterfrog Rana esculenta (RL‐genotype) is a natural hybrid between R. ridibunda (RR) and R. lessonae (LL) and reproduces by hybridogenesis, i.e. it eliminates the L‐genome from the germline and produces gametes only containing the clonally transmitted R‐genome. Because of the lack of recombination, R‐genomes are prone to accumulate spontaneous deleterious mutations. The homozygous effects of such mutations become evident in matings between hybrids: their offspring possess two clonal R‐genomes and are generally inviable. However, the evolutionary fate of R. esculenta mainly depends on the heterozygous effects of mutations on the R‐genome. These effects may be hidden in the hybrid R. esculenta because it has been shown to benefit from spontaneous heterosis. To uncouple clonal inheritance from hybridity, I crossed R. esculenta with R. ridibunda to produce nonhybrid offspring with one clonal and one sexual R‐genome, and compared their survival and larval performance with normal, sexually produced R. ridibunda tadpoles. Because environmental stress can enhance the negative effects of mutation accumulation, I measured the performance at high and low food levels. There was no indication that tadpoles with a clonal genome performed worse at either food level, suggesting that at least in the larval stage, R. esculenta benefits from heterosis without incurring any costs because of heterozygous effects of deleterious mutations on the clonally transmitted R‐genome.     

Asymmetric competition in larval amphibian communities: conservation implications for the northern crawfish frog, Rana areolata circulosa

Asymmetric competition in larval amphibians can influence population dynamics and community structure. This density-dependent regulatory mechanism may be of particular importance for rare or endangered species such as the northern crawfish frog, Rana areolata circulosa. Interspecific competition of R. areolata with two congenerics, R. blairi and R. sphenocephala, was examined in artificial ponds. Analysis of covariance (differential mortality covariate) indicated that interspecific competition increased larval period length and decreased metamorphic body mass of R. areolata. The number of metamorphs produced was lower for R. blairi ponds when reared with R. areolata at high density. Body mass at metamorphosis was larger for R. sphenocephala when reared with R. areolata, suggesting that R. areolata facilitates larval growth in R. sphenocephala. These results indicate that the larval performance of R. areolata was reduced in the presence of interspecific competitors. Although many conservation efforts emphasize the preservation of critical habitat or particular rare species, interactive effects of biotic components in the focal community may also be important demographic regulators. Received: 11 December 1997 / Accepted: 15 April 1998