Sexual dimorphisms in body size and head size are common among lizards and are often related to sexual selection on male fighting capacity (organismal performance) and territory defence. However, whether this is generally true or restricted to lizards remains untested. Here we provide data on body and head size, bite performance and indicators of mating success in the tuatara (Sphenodon punctatus), the closest living relative to squamates, to explore the generality of these patterns. First, we test whether male and female tuatara are dimorphic in head dimensions and bite force, independent of body size. Next, we explore which traits best predict bite force capacity in males and females. Finally, we test whether male bite force is correlated with male mating success in a free-ranging population of tuatara (Sphenodon punctatus). Our data confirm that tuatara are indeed dimorphic in head shape, with males having bigger heads and higher bite forces than females. Across all individuals, head length and the jaw closing in-lever are the best predictors of bite force. In addition, our data show that males that are mated have higher absolute but not relative bite forces. Bite force was also significantly correlated to condition in males but not females. Whereas these data suggest that bite force may be under sexual selection in tuatara, they also indicate that body size may be the key trait under selection in contrast to what is observed in squamates that defend territories or resources by biting.
Carry-over effects influence trait responses in later life stages as a result of early experience with environmental cues. Predation risk is an influential stressor and selection exists for early recognition of threats. In particular, invasive species may benefit from carry-over effects by preemptively recognizing and responding to novel predators via latent developmental changes and embryonic learning. In a factorial experiment, we conditioned invasive American bullfrog embryos (Lithobates catesbeianus) to the odor of a novel fish predator, largemouth bass (Micropterus salmoides) alone or in combination with injured conspecific cues. We quantified developmental carryover in the larval life stage and found that individuals conditioned to the highest risk (fish and injured conspecific cues) grew into longer bodied larvae relative to larvae from lower risk treatments. We also assessed embryonic learning, a behavioral carry-over effect, and found an interaction between embryonic conditioning and larval exposure. Behavioral responses were only found in scenarios when predation risk varied in intensity across life history stages, thus requiring a more flexible antipredator strategy. This indicates a potential trade-off between the two strategies in larval growth and development rates, and time until metamorphosis. Our results suggest that early predator exposure and carry-over effects have significant impacts on life history trajectories for American bullfrogs. This research contributes to our understanding of a potentially important invasion mechanism in an anuran species of conservation concern.
Changing patterns of precipitation and drought will dramatically influence the distribution and persistence of lentic habitats. Pond-breeding amphibians can often respond to changes in habitat by plastically shifting behavioral and developmental trait response. However, fitness tradeoffs inherent in life history strategies can carry over to impact development, behavior, and fitness in later life stages. In this experiment, we investigated carryover effects of hydroperiod permanence on the movement behavior of newly-metamorphosed juvenile Northern Red-legged Frogs (Rana aurora). Frogs were raised through metamorphosis in mesocosms under either permanent or ephemeral hydroperiod conditions. After metamorphosis, individuals were removed from the mesocosms, measured, uniquely tagged with elastomer, and moved to holding terrariums. Movement behavior was quantified under two terrain conditions: a physiologically-taxing, dry runway treatment, or a control, moist runway treatment. Individuals were given 30 min to move down the 1 × 20m enclosed structure before distance was measured. We applied a hurdle model to examine two distinct components of movement behavior: (1) the probability of moving away from the start location, and (2) movement distance. We found that hydroperiod condition had an indirect carryover effect on movement via the relationship between individual size and the propensity to move. Individuals from ephemeral mesocosm conditions metamorphosed at a smaller size but showed increased growth rates as compared to individuals from permanent hydroperiod conditions. Individual snout-vent length and runway condition (moist or dry) were significant predictors of both aspects of movement behavior. Larger individuals were more likely to move down the runway and able to move a farther distance than smaller individuals. In addition to the influence of size, dry runway conditions reduced the probability of individuals moving from the start location, but increased the distance traveled relative to the moist runway. The demonstrated cumulative impact of stressors suggests the importance of addressing direct, indirect, and carryover effects of stressors throughout ontogeny.
The development of antipredator traits is dependent on the frequency and intensity of predator exposure over evolutionary and ecological time. We hypothesized that prey species would respond with increasing accuracy when exposed to predators across generational, ontogenetic, and immediate time scales. We assessed larval Pacific chorus frog (PSRE; Pseudacris regilla) individuals that varied in population sympatry, embryonic conditioning, and immediate exposure to stocked populations of rainbow trout (Oncorhynchus mykiss). Using PSRE populations from sites with and without resident rainbow trout, we conditioned embryos to trout odor, PSRE alarm cues, trout odor in combination with alarm cues, or control water. After being hatched and reared in control water, individuals were exposed to the four predator cue treatments using a fully factorial design. Tadpoles from populations with resident rainbow trout did not behave or develop differently than tadpoles originating from fishless sites. However, we found evidence that PSRE reduced predation risk with a combination of carry‐over effect (i.e., transfer of information across life history stages) and within‐life stage phenotypically plastic mechanisms. We found both developmental and behavioral carry‐over effects: tadpoles conditioned with trout odor as embryos grew more slowly and took refuge more often than control animals. Within‐life‐stage behavioral plasticity was observed in tadpoles from all treatment groups, responding to predator cues with increased refuge use. Potentially additive effects of predator exposure on prey response should be considered when predicting the ability of prey to recognize novel threats.
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