Amanda C. Niehaus

Predicting the Physiological Performance of Ectotherms in Fluctuating Thermal Environments

SUMMARY Physiological ecologists have long sought to understand the plasticity of organisms in environments that vary widely among years, seasons and even hours. This is now even more important because human-induced climate change is predicted to affect both the mean and variability of the thermal environment. Although environmental change occurs ubiquitously, relatively few researchers have studied the effects of fluctuating environments on the performance of developing organisms. Even fewer have tried to validate a framework for predicting performance in fluctuating environments. Here, we determined whether reaction norms based on performance at constant temperatures (18, 22, 26, 30 and 34°C) could be used to predict embryonic and larval performance of anurans at fluctuating temperatures (18–28°C and 18–34°C). Based on existing theory, we generated hypotheses about the effects of stress and acclimation on the predictability of performance in variable environments. Our empirical models poorly predicted the performance of striped marsh frogs (Limnodynastes peronii) at fluctuating temperatures, suggesting that extrapolation from studies conducted under artificial thermal conditions would lead to erroneous conclusions. During the majority of ontogenetic stages, growth and development in variable environments proceeded more rapidly than expected, suggesting that acute exposures to extreme temperatures enable greater performance than do chronic exposures. Consistent with theory, we predicted performance more accurately for the less variable thermal environment. Our results underscore the need to measure physiological performance under naturalistic thermal conditions when testing hypotheses about thermal plasticity or when parameterizing models of life-history evolution.

Urban Physiology: City Ants Possess High Heat Tolerance

Urbanization has caused regional increases in temperature that exceed those measured on a global scale, leading to urban heat islands as much as 12°C hotter than their surroundings. Optimality models predict ectotherms in urban areas should tolerate heat better and cold worse than ectotherms in rural areas. We tested these predications by measuring heat and cold tolerances of leaf-cutter ants from South Americas largest city (São Paulo, Brazil). Specifically, we compared thermal tolerances of ants from inside and outside of the city. Knock-down resistance and chill-coma recovery were used as indicators of heat and cold tolerances, respectively. Ants from within the city took 20% longer to lose mobility at 42°C than ants from outside the city. Interestingly, greater heat tolerance came at no obvious expense of cold tolerance; hence, our observations only partially support current theory. Our results indicate that thermal tolerances of some organisms can respond to rapid changes in climate. Predictive models should account for acclimatory and evolutionary responses during climate change.

Striped marsh frog (Limnodynastes peronii) tadpoles do not acclimate metabolic performance to thermal variability

SUMMARY Human-induced climate change is predicted to affect not only the mean temperature of the environment but also the variability and frequency of extreme climatic events. Variability in an organisms developmental environment has the potential to markedly affect an individuals growth trajectory and physiological function, leading to impacts on individual fitness and population dynamics. Thus, it is important to consider the consequences of thermal variability on developing organisms and understand their capacity to respond to such increased variation. We investigated the capacity of larval striped marsh frogs (Limnodynastes peronii) to initiate a response to increases in the thermal variability of their developmental environment by reducing the sensitivity of their physiological rate functions to changes in temperature. In variable environments, we expected the thermal sensitivity of rate functions to decrease and their performance breadth to widen so as to buffer the effect of thermal variability. We raised larvae in stable (24°C), narrowly variable (22–26°C; mean 24°C) and widely variable (14–34°C; mean 24°C) thermal environments and measured the thermal sensitivity of their locomotor performance, heart rate, oxygen consumption and activities of two metabolic enzymes, lactate dehydrogenase and cytochrome c oxidase. We found that the temperature-dependent relationships of these physiological functions did not differ between tadpoles raised in stable or variable thermal conditions. Furthermore, the Q10 values of each response variable were virtually unaffected by treatment when measured over the entire thermal range. Our results reveal that larval amphibians exhibit little plasticity in metabolic traits to thermal variability. This lack of plasticity may have important implications for the growth and population dynamics of organisms in environments that are beginning to experience increased thermal variability.

Response of predators to Western Sandpiper nest exclosures

Abstract In 2001, predator exclosures were used to protect nests of the Western Sandpiper (Calidris mauri) in western Alaska. During the exclosure experiment, nest contents in exclosures had significantly higher daily survival rates than control nests, however, late in the study predators began to cue in on exclosures and predate the nest contents. An Arctic Fox (Alopex lagopus) dug under one exclosure and took the newly hatched chicks, and Long-tailed Jaegers (Stercorarius longicaudus) learned to associate exclosures with active nests and repeatedly visited them. The jaegers attempted to gain access to exclosed nests and pursued adult sandpipers as they emerged from the exclosures. The exclosures were removed to reduce potential mortality to adult and young sandpipers, but subsequently, post-exclosure nests had lower daily survival rates than controls during the same time period. Predation of post-exclosure eggs and chicks highlighted the lasting influence of the exclosure treatment on offspring survival because predators probably remembered nest locations. Researchers are urged to use caution when considering use of predator exclosures in areas where jaegers occur.

Females prefer athletes, males fear the disadvantaged: different signals used in female choice and male competition have varied consequences

The most commonly assumed cost for exaggerated male ornamentation is increased predation pressure owing to decreased locomotor performance or increased conspicuousness to predators. Despite its intuitive basis, the locomotor costs of male ornamentation are not well established. We tested the hypothesis that multiple male signals that are used independently during female choice and male competition could lead to varied locomotor costs. Multiple signals with varied costs could provide a more detailed indicator of overall male quality, as only the highest-quality individuals could support all costs. To test this idea, we investigated the burst locomotor consequences of multiple ornaments for males of the pacific blue-eye (Pseudomugil signifer). We evaluated five competing models relating body size, ornament size and performance traits to field measures of dominance and attractiveness. Non-propulsive male fin ornaments used during male competition were different from those used in female choice. First dorsal fin length was the most important predictor of male attractiveness, while dominance was only associated with anal fin length. Furthermore, first dorsal fin length was positively correlated with swim speed, while anal fin length was negatively associated with speed. Our study shows that multiple male signals that are used independently during male competition and female choice also differ in their associated costs. This provides a mechanism for understanding why locomotor costs for exaggerated male ornamentation are not often empirically demonstrated.

Does individual quality mask the detection of performance trade-offs? A test using analyses of human physical performance

Why are performance trade-offs so rarely detected in animals when their underlying physiological basis seems so intuitive? One possibility is that individual variation in health, fitness, nutrition, development or genetics, or ‘individual quality’, makes some individuals better or worse performers across all motor tasks. If this is the case, then correcting for individual quality should reveal functional trade-offs that might otherwise be overlooked. We tested this idea by exploring trade-offs in maximum physical performance and motor skill function in semi-professional soccer players. We assessed individual performance across five maximum ‘athletic’ tasks providing independent measures of power, stamina and speed, as well as five soccer-specific ‘motor skill’ tasks providing independent measures of foot control. We expected to find functional trade-offs between pairs of traits (e.g. endurance versus power/speed tasks or specialist–generalist performance) – but only after correcting for individual quality. Analyses of standardised raw data found positive associations among several pairs of traits, but no evidence of performance trade-offs. Indeed, peak performance across a single athletic task (degree of specialisation) was positively associated with performance averaged across all other athletic tasks (generalist). However, after accounting for an individuals overall quality, several functional trade-offs became evident. Within our quality-corrected data, 1500 m-speed (endurance) was negatively associated with squat time (power), jump distance (power) and agility speed – reflecting the expected speed–endurance trade-off; and degree of specialisation was negatively associated with average performance across tasks. Taken together, our data support the idea that individual variation in general quality can mask the detection of performance trade-offs at the whole-animal level. These results highlight the possibility that studies may spuriously conclude certain functional trade-offs are unimportant or non-existent when analyses that account for variation in general quality may reveal their cryptic presence.

How Fast Should an Animal Run When Escaping? An Optimality Model Based on the Trade-Off between Speed and Accuracy

How fast should animals move when trying to survive? Although many studies have examined how fast animals can move, the fastest speed is not always best. For example, an individual escaping from a predator must run fast enough to escape, but not so fast that it slips and falls. To explore this idea, we developed a simple mathematical model that predicts the optimal speed for an individual running from a predator along a straight beam. A beam was used as a proxy for straight-line running with severe consequences for missteps. We assumed that success, defined as reaching the end of the beam, had two broad requirements: (1) running fast enough to escape a predator, and (2) minimizing the probability of making a mistake that would compromise speed. Our model can be tailored to different systems by revising the predators maximal speed, the preys stride length and motor coordination, and the dimensions of the beam. Our model predicts that animals should run slower when the beam is narrower or when coordination is worse.

Ecological factors associated with the breeding and migratory phenology of high-latitude breeding western sandpipers

Environmental conditions influence the breeding and migratory patterns of many avian species and may have particularly dramatic effects on long-distance migrants that breed at northern latitudes. Environment, however, is only one of the ecological variables affecting avian phenology, and recent work shows that migration tactics may be strongly affected by changes in predator populations. We used long-term data from 1978 to 2000 to examine the interactions between snowmelt in western Alaska in relation to the breeding or migration phenologies of small shorebirds and their raptor predators. Although the sandpipers’ time of arrival at Alaskan breeding sites corresponded with mean snowmelt, late snowmelts did delay breeding. These delays, however, did not persist to southward migration through British Columbia, likely due to the birds’ ability to compensate for variance in the length of the breeding season. Raptor phenology at an early stopover site in British Columbia was strongly related to snowmelt, so that in years of early snowmelt falcons appeared earlier during the sandpipers’ southbound migration. These differential effects indicate that earlier snowmelt due to climate change may alter the ecological dynamics of the predator–prey system.

Measuring Edge Effects on Nest Predation in Forest Fragments: Do Finch and Quail Eggs Tell Different Stories?

Abstract Experiments assessing rates of avian nest predation often find that nests near forest edges are at high risk of predation, suggesting the importance of forest fragmentation in recent population declines of ground-nesting passerines. However, the use of quail (Coturnix spp.) eggs in nest predation experiments may confound conclusions about edge effects because only large-mouthed predators are able to consume these relatively large eggs, but both large and small-mouthed predators consume smaller passerine eggs. We directly compared predation rates on artificial nests baited with quail eggs or with zebra finch (Poephila guttata) eggs; the latter are similar in size to the eggs of many neotropical passerines. In 1998 and 1999 we placed 392 artificial ground nests at edge and interior locations in two east-central Iowa forest fragments. Predation on these nests varied with egg type (quail or finch) and location (edge or interior) and there was a significant interaction between egg type and location: predation on quail eggs was greater at edges than in the interior, whereas finch egg predation was high in both edge and interior locations. Based on tooth imprints in clay eggs, we determined that large-mouthed predators were six times more active at edges, whereas activity of small-mouthed nest predators was evenly distributed between edge and interior locations. We suggest that the use of only quail eggs can exaggerate edge effects and that finch eggs or clay eggs used in conjunction with quail eggs in artificial nests can be used to estimate relative predation rates by large- and small-mouthed predators.

Multivariate analyses of individual variation in soccer skill as a tool for talent identification and development: utilising evolutionary theory in sports science

ABSTRACT The development of a comprehensive protocol for quantifying soccer-specific skill could markedly improve both talent identification and development. Surprisingly, most protocols for talent identification in soccer still focus on the more generic athletic attributes of team sports, such as speed, strength, agility and endurance, rather than on a player’s technical skills. We used a multivariate methodology borrowed from evolutionary analyses of adaptation to develop our quantitative assessment of individual soccer-specific skill. We tested the performance of 40 individual academy-level players in eight different soccer-specific tasks across an age range of 13–18 years old. We first quantified the repeatability of each skill performance then explored the effects of age on soccer-specific skill, correlations between each of the pairs of skill tasks independent of age, and finally developed an individual metric of overall skill performance that could be easily used by coaches. All of our measured traits were highly repeatable when assessed over a short period and we found that an individual’s overall skill – as well as their performance in their best task – was strongly positively correlated with age. Most importantly, our study established a simple but comprehensive methodology for assessing skill performance in soccer players, thus allowing coaches to rapidly assess the relative abilities of their players, identify promising youths and work on eliminating skill deficits in players.