Varanid Research Articles

Anatomy of the python heart

The hearts of all snakes and lizards consist of two atria and a single incompletely divided ventricle. In general, the squamate ventricle is subdivided into three chambers: cavum arteriosum (left), cavum venosum (medial) and cavum pulmonale (right). Although a similar division also applies to the heart of pythons, this family of snakes is unique amongst snakes in having intracardiac pressure separation. Here we provide a detailed anatomical description of the cardiac structures that confer this functional division. We measured the masses and volumes of the ventricular chambers, and we describe the gross morphology based on dissections of the heart from 13 ball pythons (Python regius) and one Burmese python (P. molurus). The cavum venosum is much reduced in pythons and constitutes approximately 10% of the cavum arteriosum. We suggest that shunts will always be less than 20%, while other studies conclude up to 50%. The high-pressure cavum arteriosum accounted for approximately 75% of the total ventricular mass, and was twice as dense as the low-pressure cavum pulmonale. The reptile ventricle has a core of spongious myocardium, but the three ventricular septa that separate the pulmonary and systemic chambers--the muscular ridge, the bulbuslamelle and the vertical septum--all had layers of compact myocardium. Pythons, however, have unique pads of connective tissue on the site of pressure separation. Because the hearts of varanid lizards, which also are endowed with pressure separation, share many of these morphological specializations, we propose that intraventricular compact myocardium is an indicator of high-pressure systems and possibly pressure separation.

The cellular force‐frequency response in ventricular myocytes from the varanid lizard, Varanus exanthematicus

To investigate the cellular mechanisms underlying the negative force‐frequency relationship in the ventricle of the varanid lizard, we measured sarcomere and cell shortening, intracellular Ca2+ ([Ca2+]i), action potentials (APs) and K+ currents in isolated ventricular myocytes. Experiments were conducted between 0.2–1.0 Hz, which spans the physiological range of heart rates at 20–22°C for this species. As frequency increased, diastolic length, percent change in sarcomere length and relaxation time decreased significantly. Shortening velocity was unchanged. The rate of rise of [Ca2+]i increased, [Ca2+]i transient amplitude decreased, and diastolic [Ca2+]i increased. The first order time constant of the [Ca2+]i transient decay decreased at higher frequencies, indicating frequency dependent acceleration of relaxation, then leveled out and did not change above 0.5 Hz. The AP rate of rise was unaffected, but the AP duration (APD) decreased with increasing frequency. Peak depolarization was only significantly decreased at 1.0 Hz. The decrease in APD was not due to changes in IKr or Ito. Our results suggest that a negative FFR in varanid lizard ventricle is caused by decreased amplitude of the [Ca2+]i transient with an increase in diastolic [Ca2+]i, leading to incomplete relaxation between beats at high frequencies. This coincides with shortened action potential duration (APD) at higher frequencies.

Partitioning of temporal activity among desert lizards in relation to prey availability and temperature

AbstractPartitioning of activity time within ecological communities potentially reduces interspecific competition and increases the number of species that can coexist. We investigated temporal activity in a highly diverse lizard assemblage in the Simpson Desert, central Australia, to determine the degree of partitioning that occurs. Three periods were defined, daytime (sunrise to sunset), early night (sunset to midnight) and late night (midnight to sunrise), and live captures of lizards were tallied for each period during two sampling months (September and November 2007). We also quantified the activity times of potential invertebrate prey and measured ambient temperatures during the different time periods to investigate any associations between these factors and lizard activity. Some 77% of captures of 13 lizard species were made by day, with Ctenotus pantherinus, Egernia inornata (Scincidae) and Nephrurus levis (Gekkonidae) the only species showing extended nocturnal activity. Activity of both species of skink was recorded at temperatures 4°C lower than those for agamid and varanid lizards early in the night, and at temperatures as low as 18–20°C. Surface‐active invertebrates differed in composition between time periods and were less abundant during the late night period in the drier of the two sample months (September), but were distributed equally over time in the other month. Termites were active in subterranean galleries at night in September and mostly by day in November, but available at all times on surface/subsurface baits. We conclude that activity is distributed unevenly within this lizard assemblage, with partitioning facilitated by the ready availability of invertebrate prey and by lizards having relatively broad temperature tolerances that, in some cases, permit opportunistic exploitation of resources beyond usual times of activity.

The cellular force-frequency response in ventricular myocytes from the varanid lizard, Varanus exanthematicus

To investigate the cellular mechanisms underlying the negative force-frequency relationship (FFR) in the ventricle of the varanid lizard, Varanus exanthematicus, we measured sarcomere and cell shortening, intracellular Ca(2+) ([Ca(2+)](i)), action potentials (APs), and K(+) currents in isolated ventricular myocytes. Experiments were conducted between 0.2 and 1.0 Hz, which spans the physiological range of in vivo heart rates at 20-22 degrees C for this species. As stimulation frequency increased, diastolic length, percent change in sarcomere length, and relaxation time all decreased significantly. Shortening velocity was unaffected. These changes corresponded to a faster rate of rise of [Ca(2+)](i), a decrease in [Ca(2+)](i) transient amplitude, and a seven-fold increase in diastolic [Ca(2+)](i). The time constant for the decay of the Ca(2+) transient (tau) decreased at higher frequencies, indicating a frequency-dependent acceleration of relaxation (FDAR) but then reached a plateau at moderate frequencies and did not change above 0.5 Hz. The rate of rise of the AP was unaffected, but the AP duration (APD) decreased with increasing frequency. Peak depolarization tended to decrease, but it was only significant at 1.0 Hz. The decrease in APD was not due to frequency-dependent changes in the delayed inward rectifier (I(Kr)) or the transient outward (I(to)) current, as neither appeared to be present in varanid ventricular myocytes. Our results suggest that a negative FFR relationship in varanid lizard ventricle is caused by decreased amplitude of the Ca(2+) transient coupled with an increase in diastolic Ca(2+), which leads to incomplete relaxation between beats at high frequencies. This coincides with shortened APD at higher frequencies.

Predation on toxic cane toads (Bufo marinus) may imperil bluetongue lizards (Tiliqua scincoides intermedia, Scincidae) in tropical Australia

Context. Detecting ecological impacts of invasive species can be extremely difficult. Even major population declines may be undetectable without extensive long-term data if the affected taxon is rare and/or difficult to census, and exhibits stochastic variation in abundance as a result of other factors. Our data suggest such a situation in an iconic Australian reptile species, the bluetongue lizard. Originally restricted to Central and South America, cane toads (Bufo marinus) are rapidly spreading through tropical Australia. Most native predators have no evolutionary history of exposure to the toads’ distinctive chemical defences (bufadienolides), and many varanid lizards, elapid snakes, crocodiles and marsupials have been killed when they have attempted to consume toads. Aims. Scincid lizards have not been considered vulnerable to toad invasion; however, one lineage (the bluetongues, genus Tiliqua) consists of large omnivores that may be affected. Our field and laboratory research aimed to elucidate this concern. Methods. Nightly surveys for bluetongue lizards (Tiliqua scincoides intermedia) and cane toads were conducted along two adjacent roadways on the Adelaide River floodplain of the Northern Territory. Scent discrimination trials in the laboratory assessed lizard responses to chemical cues from three food types (native frogs, cane toads and ‘preferred foods’) by counting tongue-flicks and biting elicited by cotton swabs. A subset of lizards was presented with live toads. Key results. Numbers of bluetongues encountered during standardised field surveys in the Darwin region declined soon after toads arrived, and we have not recorded a single lizard for the last 20 months. In the laboratory, foraging responses of bluetongues were as intense to cane-toad scent as to the scent of native frogs, and many of the lizards we tested attempted to consume toads, and were poisoned as a result. Conclusions and implications. The population decline of bluetongues in this region appears to have been the direct result of fatal ingestion of toxic cane toads. Our studies thus add a scincid lizard species to the list of native Australian predators imperilled by cane-toad invasion, and point to the difficulty of detecting invader impact even for an iconic species in a system subject to detailed survey work.

Dwarf monitor lizards (Varanidae:Varanus, Odatria s. gen.) as definitive and paratenic hosts for physalopteran nematodes

Nematodes in the genus Abbreviata (Physalopteridae) were the dominant gastrointestinal helminths in the eight smallest species of dwarf varanid lizards (subgenus Odatria, Varanidae). All host species supported infection with encysted physalopterid larvae, which provoked minimal pathological changes in the host. The mean number of cysts in host tissues increased with host size across species. There were moderate prevalences of infection of adult A. hastaspicula in Varanus (O.) brevicauda (50%) and V. (O.) storri (32%), and of A. levicauda in V. (O.) pilbarensis (46%). There were no adult Abbreviata sp. infections in V. (O.) bushi. Intensity of infection was low in almost all hosts. Sympatric larger species of Varanus (subgenus Varanus) (V. gouldii, V. panoptes, and V. (O.) tristis) support heavy infections of these nematodes which may be a source of infection for dwarf varanids. The presence and species of adult Abbreviata in dwarf monitors could not be related to sympatry, host size, or host geographical range. Liability to infection is probably determined by a combination of phylogeny (in A. levicauda in V. pilbarensis and V. tristis, dwarfism being derived), and habitat (A. levicauda occurring in other arboreal species). Other facets of the physiology or ecology, not yet understood, are also probably involved. This study reveals a complex situation where evolutionary history and contemporary environments interact. Physalopteroides filicauda, Maxvachonia brygooi, Wanaristrongylus ctenoti, Hastospiculum gouldi, the cestode Oochoristica vacuolata, and a species of pentastomid were also recovered in low numbers.

Impact of Invasive Cane Toads on Australian Birds

The cane toad (Bufo marinus), a large, toxic, American anuran, was introduced to Australia in 1935. Populations of many of Australia's reptiles (snakes, varanid lizards, crocodiles) and carnivorous mammals (dasyurid marsupials) have declined because these predators are killed by the toad's powerful toxins. In contrast to these well-studied species, little is known about the cane toads impacts on Australian birds. We reviewed published and unpublished data on behavioral interactions between Australian avian predators and cane toads and collated distributional and dietary information to identify avian taxa potentially at risk from cane toad invasion. Cane toads are sympatric with 172 frog-eating bird species in Australia, and an additional 8 bird species overlap with the predicted future range of the toad. Although many bird species thus are potentially at risk, behavioral observations suggest the risk level is generally low. Despite occasional reports of Australian birds being killed when they ingest cane toads, most birds either ignore toads or survive the predation event. The apparently higher tolerance of Australian birds to toad toxins, compared with Australian reptiles and marsupials, may reflect genetic exchange between Australian birds and Asian populations that encounter other bufonid species regularly and hence have evolved the capacity to recognize or tolerate this toxic prey.

Ca2+cycling in cardiomyocytes from a high-performance reptile, the varanid lizard (Varanus exanthematicus)

The varanid lizard possesses one of the largest aerobic capacities among reptiles with maximum rates of oxygen consumption that are twice that of other lizards of comparable sizes at the same temperature. To support this aerobic capacity, the varanid heart possesses morphological adaptations that allow the generation of high heart rates and blood pressures. Specializations in excitation-contraction coupling may also contribute to the varanids superior cardiovascular performance. Therefore, we investigated the electrophysiological properties of the l-type Ca2+ channel and the Na+/Ca2+ exchanger (NCX) and the contribution of the sarcoplasmic reticulum to the intracellular Ca2+ transient (Δ[Ca2+]i) in varanid lizard ventricular myocytes. Additionally, we used confocal microscopy to visualize myocytes and make morphological measurements. Lizard ventricular myocytes were found to be spindle-shaped, lack T-tubules, and were ∼190 μm in length and 5–7 μm in width and depth. Cardiomyocytes had a small cell volume (∼2 pL), leading to a large surface area-to-volume ratio (18.5), typical of ectothermic vertebrates. The voltage sensitivity of the l-type Ca2+ channel current (ICa), steady-state activation and inactivation curves, and the time taken for recovery from inactivation were also similar to those measured in other reptiles and teleosts. However, transsarcolemmal Ca2+ influx via reverse mode Na+/Ca2+ exchange current was fourfold higher than most other ectotherms. Moreover, pharmacological inhibition of the sarcoplasmic reticulum led to a 40% reduction in the Δ[Ca2+]i amplitude, and slowed the time course of decay. In aggregate, our results suggest varanids have an enhanced capacity to transport Ca2+ through the Na+/Ca2+ exchanger, and sarcoplasmic reticulum suggesting specializations in excitation-contraction coupling may provide a means to support high cardiovascular performance.

Editorial Focus:Cardiac function of the lizard king: focus on “Ca2+cycling in cardiomyocytes from a high-performance reptile, the varanid lizard (Varanus exanthematicus)”

varanid lizards (genus Varanus), which include the Komodo dragon (Varanus komodoensis), are among the most adaptive reptile species on earth. Scientific explanation of the diverse varanid behavior (44 species spread across Australia, Asia, and Africa) relies on an understanding of their cardiac

Evolutionary relationships of sprint speed in Australian varanid lizards

AbstractEcomorphological studies often seek to link morphology and performance to relevant ecological characteristics. Varanid lizards are unique in that species can vary in body size by almost four orders of magnitude within a single genus, and a question of considerable interest is whether similar ecomorphological relationships exist when constraints on body size are reduced. We studied sprint speed in relation to size, shape and ecology for 18 species of varanid lizards. Maximal speed scaled positively with mass0.166 using least squares regression, and mass0.21 using reduced major‐axis regression. However, a curvilinear trend better described this relationship, suggesting an optimal mass of 2.83 kg with respect to speed. Including data for the komodo dragon Varanus komodoensis moves the optimum mass to 2.23 kg. We use this relationship to predict the sprint speed of the Komodo's giant extinct relative Varanus (Megalania) prisca to be 2.6–3 m s−1 similar to that of extant freshwater crocodiles Crocodylus johnstoni. When differences in speed were compared to ecological characteristics, species from open habitats were significantly faster than species from semi‐open or closed habitat types, and remained so after correction for size and phylogeny. Thus, despite large variation in body size, varanids appear to share similar associations between performance and ecology as seen in other lizard groups. Varanids did, however, differ in morphological relationships with sprint speed. Differences in relative speed were not related to relative hindlimb length, as is commonly reported for other lizard groups. Instead, size‐free forefoot length was negatively related to speed as was the size‐free thorax–abdomen length. While shorter forefeet were thought to be an adaptation to burrowing, and thus open habitats, rather than speed per se, the reduction in the thorax–abdomen length may have significant advantages to increasing speed. Biomechanical models predicting this advantage are discussed in relation to a trade‐off between speed and manoeuvrability.

Resources and energetics determined dinosaur maximal size.

Some dinosaurs reached masses that were approximately 8 times those of the largest, ecologically equivalent terrestrial mammals. The factors most responsible for setting the maximal body size of vertebrates are resource quality and quantity, as modified by the mobility of the consumer, and the vertebrate's rate of energy expenditure. If the food intake of the largest herbivorous mammals defines the maximal rate at which plant resources can be consumed in terrestrial environments and if that limit applied to dinosaurs, then the large size of sauropods occurred because they expended energy in the field at rates extrapolated from those of varanid lizards, which are approximately 22% of the rates in mammals and 3.6 times the rates of other lizards of equal size. Of 2 species having the same energy income, the species that uses the most energy for mass-independent maintenance of necessity has a smaller size. The larger mass found in some marine mammals reflects a greater resource abundance in marine environments. The presumptively low energy expenditures of dinosaurs potentially permitted Mesozoic communities to support dinosaur biomasses that were up to 5 times those found in mammalian herbivores in Africa today. The maximal size of predatory theropods was approximately 8 tons, which if it reflected the maximal capacity to consume vertebrates in terrestrial environments, corresponds in predatory mammals to a maximal mass less than a ton, which is what is observed. Some coelurosaurs may have evolved endothermy in association with the evolution of feathered insulation and a small mass.

The evolution of bipedal postures in varanoid lizards

The bipedal posture (BP) and gait of humans are unique evolutionary hallmarks, but similar stances and forms of locomotion have had enormous influences on a range of phylogenetically diverse tetrapods, particularly dinosaurs and birds, and a range of mammalian lineages, including non-human apes. The complex movements involved in bipedalism appear to have modest evolutionary origins, and it is presumed that a stable and erect posture is a prerequisite for erect strides and other bipedal movements. Facultative bipedalism in several lineages of lizards is achieved by running, but some varanid lizards (genus Varanus) exhibit BPs without running. In these cases, BPs (BPstanding) are not used as a form of locomotion; rather, BPstanding is associated with defensive displays, and such postures also probably permit better inspection of the environment. Yet, in other varanids, BPs have been observed only during combat episodes (BPcombat), where both contestants rise together and embrace in the so-called clinch phase. Numerous other species, however, show neither type of BP. Past researchers have commented that only large-bodied varanids exhibit BP, a behaviour that appears to show phylogenetic trends. We termed this idea the King–Green–Pianka (KGP) bipedal hypothesis. In this article, we address two main questions derived from the KGP hypothesis. First, what is the phylogenetic distribution of BP in Varanus and close relatives (varanoids)? Second, is BP positively correlated with the phylogenetic distribution of large body size (e.g. snout–vent length, SVL)? In addition, we asked a related question: do the lengths of the femur and tail show body size-independent adaptive trends in association with BP? Because varanid species that show BPstanding also use these postures during combat (BPcombat), both types of BP were analysed collectively and simply termed BP. Using comparative phylogenetic analyses, the reconstruction of BP required three steps, involving a single gain and two losses. Specifically, BP was widespread in the monophyletic Varanus, and the single gain occurred at the most recent common ancestor of the African clade. The two losses of BP occurred in different clades (Indo-Asian B clade and Indo-Australian Odatria clade). BPs are absent in the sister group to Varanus (Lanthanotus borneensis) and the other outgroup species (Heloderma spp.). Our phylogenetic reconstruction supports the KGP prediction that BP is restricted to large-bodied taxa. Using the Hansen model of adaptive evolution on a limited, but highly relevant morphological dataset (i.e. SVL; femur length, FL; tail length, TL), we demonstrated that these characters were not equivalent in their contribution to the evolution of BP in Varanus. SVL was significantly correlated with BP when modelled in a phylogenetic context, but the model identified random processes as dominant over adaptive evolution, suggesting that a body size threshold might be involved in the evolution of BP. A Brownian motion (BM) model outperformed the selection model in our analysis of relative TL, suggesting that TL and BP evolved independently. The selection model for relative FL outperformed the BM model, indicating that FL and BP share an adaptive history. Our non-phylogenetic analyses involving regression residuals of FL and TL vs. SVL showed no significant correlation between these characters and BP. We suggest that BP in Varanus provides a convergent or analogue model from which to investigate various forms of bipedalism in tetrapod vertebrates, especially other reptiles, such as theropod dinosaurs. Because BPstanding in varanids is possibly an incipient stage to some form of upright locomotion, its inclusion as a general model in evolutionary analyses of bipedalism of vertebrates will probably provide novel and important insights. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 97, 652–663.

Metabolic rate and endurance capacity in Australian varanid lizards (Squamata: Varanidae: Varanus)

In ecomorphological and ecophysiological studies, locomotor performance is often considered to be an intermediate step between the form of an organism and its environment. We examined this premise by measuring morphology, physiology and circular track endurance in the closely related group of Australian varanid lizards. Body size, body mass and relative body proportions were poor indicators of endurance. Body mass was not correlated with endurance and size-free lower forelimb length had only a weak relationship with endurance. Instead, maximal metabolic rate was positively correlated with endurance capacity in varanids. A comparison of varanids with other groups of lizards supported this result as varanids showed both elevated maximal metabolic rate and elevated endurance scores when compared with similar sized non-varanid lizards. There was support for a strong association between endurance with foraging mode and climate. Varanid species with higher endurance tended to be widely foraging and from xeric climates, while sit-and-wait and mesic species showed reduced endurance. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 97, 664–676.

Cellular investigation into the negative-force frequency response in ventricular myocytes from the varanid lizard

Cellular investigation into the negative-force frequency response in ventricular myocytes from the varanid lizard

Specialisations in excitation–contraction coupling contribute to high cardiovascular performance in varanid lizards

Specialisations in excitation–contraction coupling contribute to high cardiovascular performance in varanid lizards

The influence of metabolic heat production on body temperature of a small lizard, Anolis carolinensis

Little is known about the impact of increased metabolism on body temperatures of small ectotherms. We found that postprandial metabolic rates of 5 g Anolis carolinensis lizards were elevated by factorial increases of 2.3 ± 1.0 (mean ± S.E.) at 26 °C and 3.8 ± 2.1 at 30 °C over their fasting rates. Cloacal body temperatures exceeded environmental temperatures by a small amount in fasted individuals (26 °C: 0.3 ± 0.02 °C, 30 °C: 0.3 ± 0.02 °C), and by a significantly larger amount in fed individuals (26 °C: 1.0 ± 0.06 °C, 30 °C: 0.8 ± 0.08 °C). We conclude that an increased metabolic rate due to specific dynamic action leads to a small but significant elevation of body temperature in this species. Comparisons with thermal increments reported for a large (750 g) varanid lizard suggest that body size has only a minor influence on body–air temperature differentials of ectotherms. This is consistent with theoretical predictions. Finally, endogenous heat production could help elevate body temperatures in the wild and therefore play a minor role in thermoregulation.

Population‐level declines in Australian predators caused by an invasive species

AbstractThe cane toad Bufo marinus has been migrating westward across northern Australia since its introduction as a biological control agent in 1935. It has been implicated in the widespread decline of many native frog‐eating predators. To investigate the impacts of this invasive species on native predatory reptiles, annual surveys were conducted from 2001 to 2007 to document variation in the relative abundances of three varanid lizards (Varanus mertensi, Varanus mitchelli and Varanus panoptes) and one crocodile Crocodylus johnstoni species known to consume toads. In addition, the indirect effects of this variation on one agamid lizard Amphibolurus gilberti, a known prey item of V. panoptes, were also examined. Surveys were performed at two sites in northern Australia before and after the arrival of B. marinus. Significant declines in the relative abundances of all three species of varanid lizard were observed following toad arrival. Declines in the abundance of V. panoptes, V. mitchelli and V. mertensi at the two sites ranged 83–96, 71–97 and 87–93%, respectively. In contrast, A. gilberti increased by 23–26%; whereas there were no significant population‐level declines in C. johnstoni despite observations of individual effects (i.e. several dead crocodiles with B. marinus in their stomachs). These findings suggest population‐level changes in Australian lizards caused by an invasive species.

Bias in interspecific allometry: examples from morphological scaling in varanid lizards

The traditional approach to allometric analysis entails the fitting of a straight line to logarithmic transformations of the data, after which parameters in a two-parameter allometric equation are estimated by back-transformation to the original scale. We re-examined published data for dimensions of the limbs in 22 species of varanid lizards to illustrate the biases that can be introduced into allometric analyses by applying the aforementioned protocol. Statistical models fit to the original data by linear and nonlinear regression conformed better with underlying assumptions than did models obtained by back-transformation from logarithms, and the former generally were better than the latter for describing limb dimensions over the full range in body size. Allometric exponents estimated by the traditional method therefore were based on inappropriate and inaccurate statistical models and, consequently, were biased and misleading. Investigators can avoid problems such as these by performing preliminary graphical and statistical analyses on data in their original scale and by validating the fitted model. Logarithmic transformations should be used sparingly and only for cause.

The molecular evolutionary tree of lizards, snakes, and amphisbaenians

Squamate reptiles (lizards, snakes, amphisbaenians) number approximately 8200 living species and are a major component of the world's terrestrial vertebrate diversity. Recent molecular phylogenies based on protein-coding nuclear genes have challenged the classical, morphology-based concept of squamate relationships, requiring new classifications, and drawing new evolutionary and biogeographic hypotheses. Even the key and long-held concept of a dichotomy between iguanians (∼1470 sp.) and scleroglossans (all other squamates) has been refuted because molecular trees place iguanians in a highly nested position. Together with snakes and anguimorphs, iguanians form a clade – Toxicofera – characterized by the presence of toxin secreting oral glands and demonstrating a single early origin of venom in squamates. Consequently, neither the varanid lizards nor burrowing lineages such as amphisbaenians or dibamid lizards are the closest relative of snakes. The squamate timetree shows that most major groups diversified in the Jurassic and Cretaceous, 200–66 million years (Myr) ago. In contrast, five of the six families of amphisbaenians arose during the early Cenozoic, ∼60–40 Myr ago, and oceanic dispersal on floating islands apparently played a significant role in their distribution on both sides of the Atlantic Ocean. Among snakes, molecular data support the basic division between the small fossorial scolecophidians (∼370 sp.) and the alethinophidians (all other snakes, ∼2700 sp.). They show that the alethinophidians were primitively macrostomatan and that this condition was secondarily lost by burrowing lineages. The diversification of alethinophidians resulted from a mid-Cretaceous vicariant event, the separation of South America from Africa, giving rise to Amerophidia (aniliids and tropidophiids) and Afrophidia (all other alethinophidians). Finally, molecular phylogenies have made it possible to draw a detailed evolutionary history of venom among advanced snakes (Caenophidia), a key functional innovation underlying their radiation (∼2500 sp.). To cite this article: N. Vidal, S.B. Hedges, C. R. Biologies 332 (2009).

A comparison of methods for fitting allometric equations to field metabolic rates of animals

We re-examined data for field metabolic rates of varanid lizards and marsupial mammals to illustrate how different procedures for fitting the allometric equation can lead to very different estimates for the allometric coefficient and exponent. A two-parameter power function was obtained in each case by the traditional method of back-transformation from a straight line fitted to logarithms of the data. Another two-parameter power function was then generated for each data-set by non-linear regression on values in the original arithmetic scale. Allometric equations obtained by non-linear regression described the metabolic rates of all animals in the samples. Equations estimated by back-transformation from logarithms, on the other hand, described the metabolic rates of small species but not large ones. Thus, allometric equations estimated in the traditional way for field metabolic rates of varanids and marsupials do not have general importance because they do not characterize rates for species spanning the full range in body size. Logarithmic transformation of predictor and response variables creates new distributions that may enable investigators to perform statistical analyses in compliance with assumptions underlying the tests. However, statistical models fitted to transformations should not be used to estimate parameters of equations in the arithmetic domain because such equations may be seriously biased and misleading. Allometric analyses should be performed on values expressed in the original scale, if possible, because this is the scale of interest.