A phylogenetic approach to total evaporative water loss in mammals

Abstract Maintaining appropriate water balance is a constant challenge for terrestrial mammals, and this problem can be exacerbated in desiccating environments. It has been proposed that natural selection has provided desert-dwelling mammals physiological mechanisms to reduce rates of total evaporative water loss. In this study, we evaluated the relationship between total evaporative water loss and body mass in mammals by using a recent phylogenetic hypothesis. We compared total evaporative water loss in 80 species of arid-zone mammals to that in 56 species that inhabit mesic regions, ranging in size from 4 g to 3,500 kg, to test the hypothesis that mammals from arid environments have lower rates of total evaporative water loss than mammals from mesic environments once phylogeny is taken into account. We found that arid species had lower rates of total evaporative water loss than mesic species when using a dichotomous variable to describe habitat (arid or mesic). We also found that total evaporative water loss was negatively correlated with the average maximum and minimum environmental temperature as well as the maximum vapor pressure deficit of the environment. Annual precipitation and the variable Q (a measure of habitat aridity) were positively correlated with total evaporative water loss. These results support the hypothesis that desert-dwelling mammals have lower rates of total evaporative water loss than mesic species after controlling for body mass and evolutionary relatedness regardless of whether categorical or continuous variables are used to describe habitat.     

DIVERSITY AND NICHE EVOLUTION ALONG ARIDITY GRADIENTS IN NORTH AMERICAN LIZARDS (PHRYNOSOMATIDAE)

Deserts occupy approximately 12% of the Earth's land surface, and are thought to have species poor but highly specialized biotas. However, few studies have examined the evolutionary origins of desert biotas and of diversity patterns along aridity gradients. Further, it is unclear if species occurring in more extreme conditions on a given niche axis (i.e., precipitation) are more specialized for those conditions (i.e., have narrower niche breadths). We address these questions here using a time‐calibrated phylogeny and climatic data for 117 species of phrynosomatid lizards. Phrynosomatids are the most species‐rich family of lizards in North America, and are found from deserts to rainforests. Surprisingly, we find that phrynosomatids have higher richness in more arid environments. This pattern occurs seemingly because they have been present in more arid habitats longer (～55 million years), and lineages in mesic environments are recently derived from more arid‐dwelling ancestors. We find little support for the hypothesis that species in more extreme environments are more specialized. Instead, many desert‐dwelling species are broadly distributed, and species in the most mesic environments have the broadest niche breadths. In summary, phrynosomatids offer a counterexample to the idea that arid regions are inhabited by a small number of recent and highly specialized lineages.     

Water balance in desert Drosophila: lessons from non-charismatic microfauna

Water stress is a particularly important problem for insects and other small organisms in arid environments. Cactophilic fruit flies in the genus Drosophila have invaded deserts on numerous occasions, including multiple independent invasions of North American deserts. Because the evolutionary history of this genus is so well studied, we can investigate the mechanisms of adaptation in a rigorous phylogenetic context. As expected, desert fruit flies lose water less rapidly than their mesic congeners. They are also able to tolerate the loss of a greater percentage of body water, but this difference is mainly due to phylogenetic history, and does not represent an adaptation specifically to desert habitats. A laboratory analogue of desert Drosophila is provided by populations of D. melanogaster that have been subjected to selection for desiccation resistance. Selected populations resemble desert species in that they lose water slowly, relative to control populations, and are not more tolerant of dehydration stress. They differ, however, in having much higher water contents and different behavioral responses to desiccating conditions. Our comparisons of laboratory and natural populations reveal that not all possible adaptive mechanisms evolve in stressful environments. Different physiological and behavioral strategies may evolve depending upon the particular options available in the environment.     

Ecological selection pressures for C4 photosynthesis in the grasses

Grasses using the C₄ photosynthetic pathway dominate grasslands and savannahs of warm regions, and account for half of the species in this ecologically and economically important plant family. The C₄ pathway increases the potential for high rates of photosynthesis, particularly at high irradiance, and raises water-use efficiency compared with the C₃ type. It is therefore classically viewed as an adaptation to open, arid conditions. Here, we test this adaptive hypothesis using the comparative method, analysing habitat data for 117 genera of grasses, representing 15 C₄ lineages. The evidence from our three complementary analyses is consistent with the hypothesis that evolutionary selection for C₄ photosynthesis requires open environments, but we find an equal likelihood of C₄ evolutionary origins in mesic, arid and saline habitats. However, once the pathway has arisen, evolutionary transitions into arid habitats occur at higher rates in C₄ than C₃ clades. Extant C₄ genera therefore occupy a wider range of drier habitats than their C₃ counterparts because the C₄ pathway represents a pre-adaptation to arid conditions. Our analyses warn against evolutionary inferences based solely upon the high occurrence of extant C₄ species in dry habitats, and provide a novel interpretation of this classic ecological association.     

Morphological evolution in Tropidurinae squamates: an integrated view along a continuum of ecological settings

Variation in squamate foot morphology is likely relevant during evolutionary processes of habitat colonization because distinct surfaces differ in energetic and functional demands for locomotion. We combined new foot morphological data with published information of limb and tail lengths to investigate evolutionary changes possibly associated with the differential usage of ecological settings by Tropidurinae species. Several traits exhibited significant phylogenetic signal, and we performed conventional and phylogenetic regressions of PC scores (retained from Principal Components Analyses of morphometric traits) on continuous ecological indices. Tropidurines from sandy habitats exhibit larger foot soles, opposite to the evolution of narrow feet in species that use branches and rocks. Also, species that usually move along trunks present longer femora. This study provides evidence for morphological adaptations associated with substrate usage in Tropidurinae, and suggests that opposite morphological profiles might evolve associated with the use of surfaces energetically and functionally contrasting, possibly leading to trade‐offs.     

Effect of taxon sampling on recovering the phylogeny of squamate reptiles based on complete mitochondrial genome and nuclear gene sequence data

The complete nucleotide sequences of the mitochondrial (mt) genomes of three species of squamate lizards: Blanus cinereus (Amphisbaenidae), Anguis fragilis (Anguidae), and Tarentola mauritanica (Geckkonidae) were determined anew. The deduced amino acid sequences of all 13 mt protein-coding genes were combined into a single data set and phylogenetic relationships among main squamate lineages were analyzed under maximum likelihood (ML) and Bayesian Inference (BI). Within Squamata, the monophyly of Iguanidae, Anguimorpha, Amphisbaenia, Gekkota, Serpentes, and Acrodonta received high statistical support with both methods. It is particularly striking that this is the first molecular analysis that recovers the monophyly of Scincomorpha (including Scincidae, Xantusiidae, Cordylidae, and Lacertidae), although it is only supported in the Bayesian analysis, and it is sensitive to changes in the outgroup (see below). Phylogenetic relationships among the main squamate lineages could not be resolved with ML but received strong support with BI (above 95%). The newly reconstructed phylogeny of squamates does not support the Iguania-Scleroglossa split. Acrodonta and Serpentes form a clade, which is the sister group of the remaining squamate lineages. Within these, Gekkota were the first branching out, followed by Amphisbaenia, and a clade including Anguimorpha as sister group of Scincomorpha + Iguanidae. The recovered topology differed substantially from previously reported hypotheses on squamate relationships, and the relative effect of using different outgroups, genes, and taxon samplings were explored. The sister group relationship of Serpentes + Acrodonta, and their relative basal position within Squamata could be due to a long-branch attraction artifact. Phylogenetic relationships among Scincomorpha, Amphisbaenia, and Anguimorpha were found to be rather unresolved. Future improving of squamate phylogenetic relationships would rely on finding snake and acrodont species with slower mt evolutionary rates, ensuring thorough taxon coverage of squamate diversity, and incorporating more nuclear genes with appropriate evolutionary rates.     

Kidney mass and relative medullary thickness of rodents in relation to habitat, body size, and phylogeny

We tested the hypotheses that relative medullary thickness (RMT) and kidney mass are positively related to habitat aridity in rodents, after controlling for correlations with body mass. Body mass, mass-corrected kidney mass, mass-corrected RMT, mass-corrected maximum urine concentration, and habitat (scored on a semiquantitative scale of 1-4 to indicate increasing aridity) all showed statistically significant phylogenetic signal. Body mass varied significantly among habitats, with the main difference being that aquatic species are larger than those from other habitats. Mass-corrected RMT and urine concentration showed a significant positive correlation (N=38; conventional r=0.649, phylogenetically independent contrasts [IC] r=0.685), thus validating RMT as a comparative index of urine concentrating ability. RMT scaled with body mass to an exponent significantly less than 0 (N=141 species; conventional allometric slope=-0.145 [95% confidence interval (CI)=-0.172, -0.117], IC allometric slope=-0.132 [95% CI=-0.180, -0.083]). Kidney mass scaled to an exponent significantly less than unity (N=104 species; conventional slope=0.809 [95% CI=0.751, 0.868], IC slope=0.773 [95% CI=0.676, 0.871]). Both conventional and phylogenetic analysis indicated that RMT varied among habitats, with rodents from arid areas having the largest values of RMT. A phylogenetic analysis indicated that mass-corrected kidney mass was positively related to habitat aridity.     

Environmental correlates of physiological variables in marsupials

We analyzed body temperature (T(b)), basal metabolic rate (BMR), wet thermal conductance (C(wet)), and evaporative water loss (EWL) of marsupials by conventional and phylogenetically corrected regression. Allometric effects were substantial for BMR, C(wet), and EWL but not T(b). There was a strong phylogenetic signal for mass and all physiological traits. A significant phylogenetic signal remained for BMR, C(wet), and EWL even after accounting for the highly significant phylogenetic signal of mass. T(b), BMR, C(wet), and EWL allometric residuals were correlated with some diet, distribution, and climatic variables before and after correction for phylogeny. T(b) residuals were higher for marsupials from arid environments (high T(a) and more variable rainfall). The fossorial marsupial mole had a lower-than-expected T(b) residual. The allometric slope for BMR was 0.72-0.75. Residuals were consistently related to distribution aridity and rainfall variability, with species from arid and variable rainfall habitats having a low BMR, presumably to conserve energy in a low-productivity environment. The nectarivorous honey possum had a higher-than-expected BMR. For C(wet), the allometric slope was 0.55-0.62; residuals were related to diet, with folivores having low and insectivores high C(wet) residuals. The allometric slope for EWL was 0.68-0.73. EWL residuals were consistently correlated with rainfall variability, presumably facilitating maintenance of water balance during dry periods.     

Evolution of jumping capacity in Tropidurinae lizards: does habitat complexity influence obstacle‐crossing ability?

Jumping performance is relevant for lizards in many ecological contexts and might be favoured during the colonization of structurally complex habitats. Although ground-dwelling lizards use jumps to overcome small obstacles in their natural environments, jumping capacity has been mostly studied in arboreal species. Here, we analysed the evolution of jumping behaviour and performance in lizards from eight ground-dwelling species of Tropidurinae attempting to cross obstacles of different heights in a jumping track, both when undisturbed and under continuous stimulation. To establish ecological correlates with habitat complexity, individuals from two contrasting Brazilian habitats, the arid Caatingas (sand species) and the savannah-like Cerrados (rock species), were compared. Rock species jumped more often and crossed higher obstacles than sand ones in both tests, and performed more vertical than horizontal jumps. Although sand species performed less jumps, they were more successful at crossing the obstacles presented in comparison with rock species. Phylogenetic analyses confirmed these findings and demonstrated a large divergence in jumping capacity between sister-species from different habitats. Therefore, the differences in propensity and endurance for jumping activity appear to be independent of phylogenetic relationships in Tropidurinae and likely reflect an adaptation to the contrasting environments inhabited. The ecological implications of these findings are discussed.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 393–402.     

Dispersal patterns of Squamata in the Mediterranean: An evolutionary perspective

Mediterranean islands have a high diversity of squamates, although they are unevenly distributed. This variability in the composition of the reptile assemblages across islands may have been influenced by differences in the colonization abilities of these species. To evaluate the dispersal capacities of squamate species, we modeled their sea routes using cost surface models. We estimated the effects of some life‐history traits and the phylogenetic signal in the characteristics of the modeled dispersal paths. We hypothesized that a significant phylogenetic signal should be present if the dispersal ability is enhanced by traits shared among evolutionarily related species. The results showed that no phylogenetic signal was present in the characteristics of the dispersal paths (i.e., in the distance traveled/bypassed sea depth). Thus, no superior island‐colonizer lineages were detected in Mediterranean Squamata. However, our analyses also revealed that small‐sized lizards were superior to other groups of squamates at dispersing over long distances on the sea.     

Out of Africa: biogeographic history of the open‐habitat chats (Aves,Muscicapidae: Saxicolinae) across arid areas of the old world

Arid and semi‐arid areas constitute a prominent feature of the earth today, especially in Asia and Africa. Their formation started in the middle Miocene with increased stepwise aridification since the Pliocene. This aridification had strong ecological and evolutionary consequences and not only led to fragmentation of moist‐adapted biota, but also fostered the evolution of arid‐adapted taxa from mesic ancestors and triggered speciation within arid areas. The open‐habitat chats, a clade within Saxicolinae (Aves, Muscicapidae), constitute one of the most significant arid‐adapted passerine groups of Africa and Eurasia. Here, we present a temporal and spatial framework for the diversification of open‐habitat chats, using probabilistic approaches for the reconstruction of their biogeographic history based on a time‐calibrated multilocus molecular phylogenetic hypothesis. The diversification of open‐habitat chats was initiated in the late Miocene at around 7.4 Ma, most likely in sub‐Saharan Africa. Southern Africa and the Horn of Africa acted as centres of diversification and biogeographic expansion. From the latter area, the Arabo‐Sindic region and subsequently further parts of Eurasia and North Africa were colonized. The colonization history out of sub‐Saharan Africa contrasts with that of several other songbird clades, where a biogeographic expansion from Eurasia or northern Africa to southern Africa was prevalent. Habitat fragmentation through forest expansions during intermittent wetter periods in Africa influenced diversification in several clades. However, phases of increased aridity, with hyperarid regions acting as drivers of vicariance, seem to have also been important in radiations of the Arabo‐Sindic region and the Horn of Africa during the Pleistocene. Different processes such as colonization of new areas followed by vicariance or speciation across ecotones might have played a role throughout the radiation of open‐habitat chats.     

Thermal biology of African lovebirds and Australian grass parakeets

We compared aspects of the thermal biology of two groups of small parrots, of similar body mass, each derived from a range of habitat types, varying in aridity, but indigenous to either southern Africa or Australia. By accounting for phylogenetic differences, we were able to question whether arid zone species have lower metabolic rates and greater thermal tolerances than mesic species in relation to the “pre-adapted” and “post-arrival adaptation” hypotheses. Four species of African lovebird (Agapornis) and four species of Australian grass parakeet (one Neopsephotus and three Neophema species) were investigated. The Rosy-faced Lovebird (Agapornis roseicollis), Bourke's Parakeet (Neopsephotus bourkii) and the Scarlet-chested Parakeet (Neophema splendida) were categorised as arid zone species, Fischer's Lovebird (Agapornis fischeri), the Black-masked Lovebird (Agapornis personatus) and the Elegant Parakeet (Neophema elegans) as semi-arid zone species, and the Black-cheeked Lovebird (Agapornis nigrigenis) and the Turquoise Parakeet (Neophema pulchella) as mesic zone species. Conventional and phylogenetically independent statistical methods yielded no significant differences in the basal metabolic rates of birds from different habitats or between the species assemblages from Africa and Australia.     

Repeated evolution of amphibious behavior in fish and its implications for the colonization of novel environments

We know little about on how frequently transitions into new habitats occur, especially the colonization of novel environments that are the most likely to instigate adaptive evolution. One of the most extreme ecological transitions has been the shift in habitat associated with the move from water to land by amphibious fish. We provide the first phylogenetic investigation of these transitions for living fish. Thirty‐three families have species reported to be amphibious and these are likely independent evolutionary origins of fish emerging onto land. Phylogenetic reconstructions of closely related taxa within one of these families, the Blenniidae, inferred as many as seven convergences on a highly amphibious lifestyle. Taken together, there appear to be few constraints on fish emerging onto land given amphibious behavior has evolved repeatedly many times across ecologically diverse families. The colonization of novel habitats by other taxa resulting in less dramatic changes in environment should be equally, if not, more frequent in nature, providing an important prerequisite for subsequent adaptive differentiation.     

Molecular phylogeny of the mycorrhizal desert truffles (Terfezia and Tirmania), host specificity and edaphic tolerance

Terfezia and Tirmania, so called desert truffles, are mycorrhizal fungi mostly endemic to arid and semi-arid areas of the Mediterranean Region, where they are associated with Helianthemum species. The aim of this work was to study the phylogenetic relationships in these pezizalean hypogeous fungi. The restriction fragment length polymorphism (RFLP) and DNA sequences of internal transcribed spacers (ITS) of the nuclear rDNA were studied for several morphological species, Terfezia arenaria, T. boudieri, T. claveryi, T. leptoderma, T. terfezioides (=Mattirolomyces terfezioides), Tirmania nivea and T. pinoyi. The sequences were analyzed with distance and parsimony methods. Phylogenetic analyses indicated a close genetic relationship between Tirmania and Terfezia. They may have arisen from a single evolutionary lineage of pezizalean fungi that developed the hypogeous habit as an adaptation to heat and drought in Mediterranean ecosystems. This analysis also supports the re-establishment of the genus Mattirolomyces. The genera Tirmania and Terfezia were monophyletic, and morphological species corresponded to phylogenetic species. The Tirmania clade comprises desert truffles with smooth spores and amyloid asci, which were found in deserts. The Terfezia clade grouped species found in semi-arid habitats having ornamented and spherical spores. These species are adapted to exploit different types of soil (either acid or basic soils) in association with specific hosts (either basophilous or acidophilous species). Although other factors might also play a role, host specialization and edaphic tolerances (fungus and/or host tolerances) might be the key in the species diversity of these genera.     

A multilocus perspective on colonization accompanied by selection and gene flow

The colonization of novel habitats involves complex interactions between founder events, selection, and ongoing migration, and can lead to diverse evolutionary outcomes from local extinction to adaptation to speciation. Although there have been several studies of the demography of colonization of remote habitats, less is known about the demographic consequences of colonization of novel habitats within a continuous species range. Populations of the Eastern Fence Lizard, Sceloporus undulatus, are continuously distributed across two dramatic transitions in substrate color in southern New Mexico and have undergone rapid adaptation following colonization of these novel environments. Blanched forms inhabit the gypsum sand dunes of White Sands and melanic forms are found on the black basalt rocks of the Carrizozo lava flow. Each of these habitats formed within the last 10,000 years, allowing comparison of genetic signatures of population history for two independent colonizations from the same source population. We present evidence on phenotypic variation in lizard color, environmental variation in substrate color, and sequence variation for mitochondrial DNA and 19 independent nuclear loci. To confirm the influence of natural selection and gene flow in this system, we show that phenotypic variation is best explained by environmental variation and that neutral genetic variation is related to distance between populations, not partitioned by habitat. The historical demography of colonization was inferred using an Approximate Bayesian Computation (ABC) framework that incorporates known geological information and allows for ongoing migration with the source population. The inferences differed somewhat between mtDNA and nuclear markers, but overall provided strong evidence of historical size reductions in both white sand and black lava populations at the time of colonization. Populations in both novel habitats appear to have undergone partial but incomplete recovery from the initial bottleneck. Both ABC analyses and measures of mtDNA sequence diversity also suggested that population reductions were more severe in the black lava compared to the white sands habitat. Differences observed between habitats may be explained by differences in colonization time, habitat geometry, and strength or response to natural selection for substrate matching. Finally, effective population size reductions in this system appear to be more dramatic when colonization is accompanied by a change in selection regime. Our analyses are consistent with a demographic cost of adaptation to novel environments and show that it is possible to infer aspects of the historical demography of local adaptation even in the presence of ongoing gene flow.     

Out of the Bassian province: historical biogeography of the Australasian platycercine parrots (Aves,Psittaciformes)

Schweizer, M., Güntert, M. & Hertwig, S. T. (2012). Out of the Bassian province: historical biogeography of the Australasian platycercine parrots (Aves, Psittaciformes). —Zoologica Scripta, 42, 13–27. Aridification from mid‐Miocene onwards led to a fragmentation of mesic biomes in Australia and an expansion of arid habitats. This influenced the diversification of terrestrial organisms, and the general direction of their radiations is supposed to have been from mesic into drier habitats. We tested this hypothesis in the platycercine parrots that occur in different habitats in Australia and also colonized Pacific islands. We inferred their temporal and spatial diversification patterns using a Bayesian relaxed molecular clock approach based on three nuclear and two mitochondrial genes and model‐based biogeographic reconstructions. The Bassian biota was found to be the centre of origin of platycercine parrots and diversification within two of their three clades coincided with the beginning of aridification of Australia. The associated habitat changes may have catalysed their radiation through adaptation to arid environments and vicariance because of the fragmentation of non‐arid habitats. The small oceanic islands of Melanesia contributed as stepping stones for the colonization of New Zealand from Australia.     

Parallel adaptive radiations in arid and temperate Australia: molecular phylogeography and systematics of the Egernia whitii (Lacertilia: Scincidae) species group

It has been an enduring belief that increasing aridity combined with Pliocene‐Pleistocene glacial cycles resulted in the formation of distinct arid zone and temperate zone faunas within Australia. We assembled a molecular phylogeny for the Egernia whitii species group, an endemic group of skinks that comprises representatives from arid and temperate Australia, in order to test several biogeographical hypotheses regarding the origin of the Australian arid zone fauna. Sequence data were obtained from ten of the 11 species within the species group, plus three other Egernia species and an outgroup (Eulamprus heatwolei). We targeted portions of the ND4 (696 base pairs) and 16S rRNA (500 bp) mitochondrial genes and the β‐Fibrinogen 7th Intron nuclear gene (648 bp). The edited alignment comprised 1844 characters, of which 551 (30%) were variable and 382 (69%) were parsimony informative. We analysed the data using maximum likelihood and Bayesian techniques and produced a single optimal tree. Our phylogeny strongly supports two major clades within the species group, corresponding to temperate‐adapted rock‐dwelling species and arid‐adapted obligate burrowing species. However, the phylogenetic affinities of E. pulchra were not resolved. Our topology indicates that the New South Wales population of E. margaretae is actually E. whitii and reveals that E. margaretae margaretae and E. m. personata are distinct species. There also appears to be a major phylogeographical break within E. whitii occurring in eastern Victoria. Although our data supported several previously proposed phylogenetic relationships, Shimodaira–Hasegawa tests soundly rejected several suggested affinities between certain species. The arid zone members of the E. whitii species group had been suggested to have originated as a result of multiple periods of colonization during the Pleistocene glaciation cycles. However, our genetic data suggest a single origin (presumably from a semiarid E. multiscutata‐like ancestor) for the arid zone members of the group prior to the Plio‐Pleistocene, probably during the late Miocene to early Pliocene. Our topology displays substantial sequence divergence between species with short internodes and long terminal branches, indicating rapid adaptive radiations within the arid and temperate zones. The presence of temperate‐adapted species within more mesic refugia of the arid zone suggests that the necessary adaptations to aridity for colonizing the dry interior of the continent have not evolved since the initial period of adaptive radiation, despite the long evolutionary history of the species group. © 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 83 , 157–173.     

The evolutionary history of Melianthus (Melianthaceae)

The evolutionary origins of the morphological and taxonomic diversity of angiosperms is poorly known. We used the genus Melianthus to explore the diversification of the southern African flora. Melianthus comprises eight species, and a phylogeny based on one nuclear and two plastid genes, as well as a morphological data set, confirmed that the genus is monophyletic. The two earliest diverging lineages are found in relatively mesic habitats, whereas the two terminal clades (an eastern and a western clade), each with three species, favor more arid habitats. The eastern clade is largely restricted to the summer-rainfall parts of southern Africa, and the western clade is found in winter-rainfall region. Molecular dating indicates a mid-Tertiary origin of the genus, with diversification of the eastern and western clades coincident with the Late Miocene-Pliocene uplift of the Escarpment mountains and the establishment of summer aridity along the west coast. The remarkably complex flowers are indicative of sunbird pollination, but many smaller birds can also visit. Speciation may be the consequence of allopatric divergence into edaphic-climatic niches. Divergence in flower and inflorescence morphology might be in response to the divergent pressures for nectar conservation in arid regions coupled with the need for signaling to avian pollinators in generally shrubby vegetation.     

A phylogenetic test for adaptive convergence in rock-dwelling lizards

Phenotypic similarity of species occupying similar habitats has long been taken as strong evidence of adaptation, but this approach implicitly assumes that similarity is evolutionarily derived. However, even derived similarities may not represent convergent adaptation if the similarities did not evolve as a result of the same selection pressures; an alternative possibility is that the similar features evolved for different reasons, but subsequently allowed the species to occupy the same habitat, in which case the convergent evolution of the same feature by species occupying similar habitats would be the result of exaptation. Many lizard lineages have evolved to occupy vertical rock surfaces, a habitat that places strong functional and ecological demands on lizards. We examined four clades in which species that use vertical rock surfaces exhibit long hindlimbs and flattened bodies. Morphological change on the phylogenetic branches leading to the rock-dwelling species in the four clades differed from change on other branches of the phylogeny; evolutionary transitions to rock-dwelling generally were associated with increases in limb length and decreases in head depth. Examination of particular characters revealed several different patterns of evolutionary change. Rock-dwelling lizards exhibited similarities in head depth as a result of both adaptation and exaptation. Moreover, even though rock-dwelling species generally had longer limbs than their close relatives, clade-level differences in limb length led to an overall lack of difference between rock- and non-rock-dwelling lizards. These results indicate that evolutionary change in the same direction in independent lineages does not necessarily produce convergence, and that the existence of similar advantageous structures among species independently occupying the same environment may not indicate adaptation.     

Ecological and evolutionary physiology of desert birds: a progress report

The adaptive significance of mechanisms of energy and water conservation among species of desert rodents, which avoid temperature extremes by remaining within a burrow during the day, is well established. Conventional wisdom holds that arid-zone birds, diurnal organisms that endure the brunt of their environment, occupy these desert climates because of the possession of physiological design features common to all within the class Aves. We review studies that show that desert birds may have evolved specific features to deal with hot desert conditions including: a reduced basal metabolic rate (BMR) and field metabolic rate (FMR), and lower total evaporative water loss (TEWL) and water turnover (WTO).Previous work on the comparative physiology of desert birds relied primarily on information gathered on species from the deserts of the southwestern U.S., which are semi-arid habitats of recent geologic origin. We include data on species from Old World deserts, which are geologically older than those in the New World, and place physiological responses along an aridity axis that includes mesic, semi-arid, arid, and hyperarid environments.The physiological differences between desert and mesic birds that we have identified using the comparative method could arise as a result of acclimation to different environments, of genetic change mediated by selection, or both. We present data on the flexibility of BMR and TEWL in Hoopoe Larks that suggest that phenotypic adjustments in these variables can be substantial. Finally, we suggest that linkages between the physiology of individual organism and its life-history are fundamental to the understanding of life-history evolution.