Phylogenetic analysis of adaptation in comparative physiology and biomechanics: overview and a case study of thermal physiology in treefrogs.

Comparative phylogenetic studies of adaptation are uncommon in biomechanics and physiology. Such studies require data collection from many species, a challenge when this is experimentally intensive. Moreover, researchers struggle to employ the most biologically appropriate phylogenetic tools for identifying adaptive evolution. Here, we detail an established but greatly underutilized phylogenetic comparative framework - the Ornstein-Uhlenbeck process - that explicitly models long-term adaptation. We discuss challenges in implementing and interpreting the model, and we outline potential solutions. We demonstrate use of the model through studying the evolution of thermal physiology in treefrogs. Frogs of the family Hylidae have twice colonized the temperate zone from the tropics, and such colonization likely involved a fundamental change in physiology due to colder and more seasonal temperatures. However, which traits changed to allow colonization is unclear. We measured cold tolerance and characterized thermal performance curves in jumping for 12 species of treefrogs distributed from the Neotropics to temperate North America. We then conducted phylogenetic comparative analyses to examine how tolerances and performance curves evolved and to test whether that evolution was adaptive. We found that tolerance to low temperatures increased with the transition to the temperate zone. In contrast, jumping well at colder temperatures was unrelated to biogeography and thus did not adapt during dispersal. Overall, our study shows how comparative phylogenetic methods can be leveraged in biomechanics and physiology to test the evolutionary drivers of variation among species.

The interacting effects of ungulate hoofprints and predatory native ants on metamorph cane toads in tropical Australia.

Many invasive species exploit the disturbed habitats created by human activities. Understanding the effects of habitat disturbance on invasion success, and how disturbance interacts with other factors (such as biotic resistance to the invaders from the native fauna) may suggest new ways to reduce invader viability. In tropical Australia, commercial livestock production can facilitate invasion by the cane toad (Rhinella marina), because hoofprints left by cattle and horses around waterbody margins provide distinctive (cool, moist) microhabitats; nevertheless the same microhabitat can inhibit the success of cane toads by increasing the risks of predation or drowning. Metamorph cane toads actively select hoofprints as retreat-sites to escape dangerous thermal and hydric conditions in the surrounding landscape. However, hoofprint geometry is important: in hoofprints with steep sides the young toads are more likely to be attacked by predatory ants (Iridomyrmex reburrus) and are more likely to drown following heavy rain. Thus, anthropogenic changes to the landscape interact with predation by native taxa to affect the ability of cane toads in this vulnerable life-history stage to thrive in the harsh abiotic conditions of tropical Australia.

Larger body size at metamorphosis enhances survival, growth and performance of young cane toads (Rhinella marina).

Body size at metamorphosis is a key trait in species (such as many anurans) with biphasic life-histories. Experimental studies have shown that metamorph size is highly plastic, depending upon larval density and environmental conditions (e.g. temperature, food supply, water quality, chemical cues from conspecifics, predators and competitors). To test the hypothesis that this developmental plasticity is adaptive, or to determine if inducing plasticity can be used to control an invasive species, we need to know whether or not a metamorphosing anuran's body size influences its subsequent viability. For logistical reasons, there are few data on this topic under field conditions. We studied cane toads (Rhinella marina) within their invasive Australian range. Metamorph body size is highly plastic in this species, and our laboratory studies showed that larger metamorphs had better locomotor performance (both on land and in the water), and were more adept at catching and consuming prey. In mark-recapture trials in outdoor enclosures, larger body size enhanced metamorph survival and growth rate under some seasonal conditions. Larger metamorphs maintained their size advantage over smaller siblings for at least a month. Our data support the critical but rarely-tested assumption that all else being equal, larger body size at metamorphosis is likely to enhance an individual's long term viability. Thus, manipulations to reduce body size at metamorphosis in cane toads may help to reduce the ecological impact of this invasive species.

Helminth parasites of the leopard frog Lithobates sp. Colima (Amphibia: Ranidae) from Colima, Mexico.

The helminth fauna inhabiting Lithobates sp. Colima from Ticuizitán, Colima, Mexico, comprises 10 species: 4 digeneans ( Clinostomum sp., Glypthelmins quieta , Haematoloechus sp., and Langeronia macrocirra ), 5 nematodes ( Aplectana itzocanensis , Cosmocerca podicipinus , Foleyellides striatus , Oswaldocruzia subauricularis , and Rhabdias sp.), and 1 cestode (Cyclophyllidea). Glypthelmins quieta , L. macrocirra , and A. itzocanensis represent new host records. These observations, added to previous records from Acapulco, Guerrero, Mexico, indicate that the helminth fauna of Lithobates sp. from Colima comprises 25 taxa. Frogs are being parasitized by 3 infection routes: ingestion of intermediate host, skin penetration by larval forms, and transmission by vectors. Species of Aplectana , Cosmocerca , Foleyellides , and Oswaldocruzia occurred in high prevalence in Colima, similar to a previous study on the same frog species from Guerrero. In Colima, Glypthelmins , Haematoloechus , and Rhabdias also occurred in high prevalence. Haematoloechus species reached the highest mean intensity in both localities. The semiaquatic habits of this species of frog and the availability of particular feeding resources appear to determine the helminth composition and infection levels; however, co-speciation events also play an important role structuring these helminth communities.