Interactions between locomotion,feeding, and bodily elongation during the evolution of snakes

Information from lizard lineages that have evolved a highly elongate (snake‐like) body form may clarify the selective forces important in the early evolution of snakes. Lizards have evolved bodily elongation via two distinct routes: as an adaptation to burrowing underground or to rapid locomotion above ground. These two routes involve diametrically opposite modifications to the body plan. Burrowing lizards have elongate trunks, small heads, short tails, and relatively constant body widths, whereas surface‐active taxa typically have shorter trunks, wider heads, longer tails, and more variable body widths. Snakes resemble burrowing rather than surface‐active (or aquatic) lizards in these respects, suggesting that snakes evolved from burrowing lizards. The trunk elongation of burrowing lizards increases the volume of the alimentary tract, so that an ability to ingest large meals (albeit consisting of small individual prey items) was present in the earliest snakes. Subsequent shifts to ingestion of wide‐bodied prey came later, after selection dismantled other gape‐constraining morphological attributes, some of which may also have arisen as adaptations to burrowing through hard soil (e.g. relatively small heads, rigid skulls). Adaptations of snake skulls to facilitate ingestion of large prey have evolved to compensate for the reduction of relative head size accompanying bodily elongation; relative to predator body mass, maximum sizes of prey taken by snakes may not be much larger than those of many lizards. This adaptive scenario suggests novel functional links between traits, and a series of testable predictions about the relationships between squamate morphology, habitat, and trophic ecology. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 293–304.     

A neglected life-history trait: clutch-size variance in snakes

Most analyses of life-history traits have focused on mean values rather than their associated variance. We review published and original data on snakes, including records gathered over many years on single populations, to examine patterns in clutch-size variability in these animals. Within single populations, the coefficient of variation of clutch size did not vary significantly with maternal body size, or among years. The stability of clutch-size variance through time is consistent with experimental studies showing no significant influence of food intake rates on this characteristic. Clutch-size variances did not differ between viviparous and oviparous snakes, but were dependent upon allometric relationships involving maternal body size and the relationship between clutch size and body size. Clutch-size variability was highest in species with relatively variable female sizes, and with a high rate of increase in clutch size with increasing body size. These two factors acted to magnify the extent of clutch-size variability engendered by variability in maternal body sizes. The relationships among these variables were similar in the two squamate Suborders, but the larger body sizes and mean clutch sizes of snakes resulted in clutch-size variances being higher in snakes than in lizards.     

Sexual dimorphism, reproductive biology, and food habits of two species of African filesnakes (Mehelya, Colubridae)

The ecology and general biology of African snakes remains virtually unstudied, even in highly distinctive species such as the filesnakes (genera Mehelya and Gonionotophis ). Our measurements and dissections of preserved specimens provided information on body sizes, sexual dimorphism in size and bodily proportions, clutch sizes, and food habits of two Mehelya species. In both M. capensis and M. nyassae , females attain sexual maturity at the same size as conspecific males, but grow to much larger sizes. Mehelya capensis displays extreme differences in body shape between males and females at the same body length: females have longer and wider heads, thicker bodies, and larger eyes (relative to both head length and head width) than do conspecific males. Dimorphism in body proportions is less marked in M. nyassae. Female reproductive cycles are seasonal in M. capensis , and clutch sizes are larger in this species than in its smaller congener (5-11 eggs in M. capensis , 2-6 eggs in M. nyassae ).
Contrary to popular wisdom, Mehelya are not specialized ophiophages. Mehelya nyassae feeds primarily upon lygosomatine skinks, including many fossorial taxa. Mehelya capensis has a broader diet, feeding on a wide variety of terrestrial lizards (especially agamids and gerrhosaurids) and snakes. Toads are also common prey items. The diversity of prey types taken by M. capensis suggests that these snakes may use ambush predation as well as active foraging. Mehelya is strongly convergent with Asian elapids of the genus Bungarus in its morphology (triangular body shape; powerful jaws; visible interstitial skin), behaviour (nocturnality; reluctance to bite when harassed), and diet (feeding on elongate reptiles, including snakes). Observations of preyhandling and ingestion by captive snakes are needed to clarify possible selective forces for the evolution of the unusual traits shared by these taxa.     

Energetic and physiological correlates of prey handling and ingestion in lizards and snakes

In this review, we summarize the energetic and physiological correlates of prey handling and ingestion in lizards and snakes. There were marked differences in the magnitude of aerobic metabolism during prey handling and ingestion between these two groups, although they show a similar pattern of variation as a function of relative prey mass. For lizards, the magnitude of aerobic metabolism during prey handling and ingestion also varied as a function of morphological specializations for a particular habitat, prey type, and behavior. For snakes, interspecific differences in aerobic metabolism during prey handling seem to be correlated with adaptations for prey capture (venom injection vs. constriction). During ingestion by snakes, differences in aerobic metabolism might be due to differences in cranial morphology, although allometric effects might be a potentially confounded effect. Anaerobic metabolism is used for prey handling and ingestion, but its relative contribution to total ATP production seems to be more pronounced in snakes than in lizards. The energetic costs of prey handling and ingestion are trivial for both groups and cannot be used to predict patterns of prey-size selection. For lizards, it seems that morphological and ecological factors set the constraints on prey handling and ingestion. For snakes, besides these two factors, the capacity of the cardio-respiratory system may also be an important factor constraining the capacity for prey handling and ingestion.     

Morphology and frictional properties of scales of Pseudopus apodus (Anguidae,Reptilia)

In the lizard family Anguidae different levels of limb reduction exist up to a completely limbless body. The locomotion patterns of limbless anguid lizards are similar to the undulating and concertina movements of snakes. Additionally, anguid lizards frequently use a third mode of locomotion, called slide-pushing. During slide-pushing the undulating moving body slides on the ground, while the posterior part of the body is pressed against the substrate. Whereas the macroscopic and microscopic adaptations of snake scales to limbless locomotion are well described, the micromorphology of anguid lizard scales has never been examined. Therefore we studied the macro- and micromorphology of the scales of Pseudopus apodus, an anguid lizard with a snakelike body. In addition, we measured the frictional properties of Pseudopus scales. Our data show that the microstructures of the ventral scales of this anguid lizard are less developed than in snakes. We found, however, a rostro-caudal gradient in macroscopic structuring. Whereas the ventral side of the anterior body was nearly unstructured, the tail had macroscopic longitudinal ridges. Our frictional measurements on rough substrates revealed that the ridges provide a frictional anisotropy: friction was higher in the lateral than in the rostral direction. The observed frictional properties are advantageous for a tail-based slide-pushing locomotion, for which a tail with a high lateral friction is most effective in generating propulsion.     

Allometry of reproduction in two species of gekkonid lizards (Gehyra): effects of body size miniaturization on clutch and egg sizes

In squamate reptiles there is an allometric pattern for small-bodied females to have smaller clutches and proportionally larger eggs than large-bodied females, and this pattern occurs both among and within species. The allometric patterns in two species of the gecko Gehyra were studied to see how evolutionary reductions in adult body size affect fecundity and offspring size among species, and how these changes affect allometric relationships within species. Gehyra dubia has two eggs per clutch (the typical clutch size for gekkonid lizards), whereas the smallerbodied G. variegata has a single egg per clutch. Within both species, egg size increased with female body size. The data are consistent with at least two mechanistic hypotheses: (1) that the width of the pelvis constrains egg size; and (2) in species with invariant clutch sizes, larger females can only allocate additional energy towards egg size and not number. More direct tests of these hypotheses are warranted. Miniaturization of body sizes in Gehyra is correlated with a clutch size reduction of 50% (from two to one), and a large (1.7-fold) compensatory increase in relative egg mass. However, the small-bodied G. variegata (one egg per clutch) had a lower relative clutch mass than did G. dubia. These findings have implications for understanding the influence of evolutionary reductions in body size on reproductive traits, and for allometric trends in squamate reptiles in general.     

Morphology,Reproduction and Diet in Australian and Papuan Death Adders (Acanthophis,Elapidae)

Death adders (genus Acanthophis) differ from most other elapid snakes, and resemble many viperid snakes, in their thickset morphology and ambush foraging mode. Although these snakes are widely distributed through Australia and Papua New Guinea, their basic biology remains poorly known. We report morphological and ecological data based upon dissection of >750 museum specimens drawn from most of the range of the genus. Female death adders grow larger than conspecific males, to about the same extent in all taxa (20% in mean adult snout-vent length,  =  SVL). Most museum specimens were adult rather than juvenile animals, and adult males outnumbered females in all taxa except A. pyrrhus. Females have shorter tails (relative to SVL) than males, and longer narrower heads (relative to head length) in some but not all species. The southern A. antarcticus is wider-bodied (relative to SVL) than the other Australian species. Fecundity of these viviparous snakes was similar among taxa (mean litter sizes 8 to 14). Death adders encompass a broad range of ecological attributes, taking a wide variety of vertebrate prey, mostly lizards (55%), frogs and mammals (each 21%; based on 217 records). Dietary composition differed among species (e.g. frogs were more common in tropical than temperate-zone species), and shifted with snake body size (endotherms were taken by larger snakes) and sex (male death adders took more lizards than did females). Overall, death adders take a broader array of prey types, including active fast-moving taxa such as endotherms and large diurnal skinks, than do most other Australian elapids of similar body sizes. Ambush foraging is the key to capturing such elusive prey.     

Do lizards and snakes really differ in their ability to take large prey? A study of relative prey mass and feeding tactics in lizards

Adaptations of snakes to overpower and ingest relatively large prey have attracted considerable research, whereas lizards generally are regarded as unable to subdue or ingest such large prey items. Our data challenge this assumption. On morphological grounds, most lizards lack the highly kinetic skulls that facilitate prey ingestion in macrostomate snakes, but (1) are capable of reducing large items into ingestible-sized pieces, and (2) have much larger heads relative to body length than do snakes. Thus, maximum ingestible prey size might be as high in some lizards as in snakes. Also, the willingness of lizards to tackle very large prey items may have been underestimated. Captive hatchling scincid lizards (Bassiana duperreyi) offered crickets of a range of relative prey masses (RPMs) attacked (and sometimes consumed parts of) crickets as large as or larger than their own body mass. RPM affected foraging responses: larger crickets were less likely to be attacked (especially on the abdomen), more likely to be avoided, and less likely to provide significant nutritional benefit to the predator. Nonetheless, lizards successfully attacked and consumed most crickets ≤35% of the predator’s own body mass, representing RPM as high as for most prey taken by snakes. Thus, although lizards lack the impressive cranial kinesis or prey-subduction adaptations of snakes, at least some lizards are capable of overpowering and ingesting prey items as large as those consumed by snakes of similar body sizes.     

Identifying sources of variation in reproductive and life-history traits among five populations of a Chinese lizard (Takydromus septentrionalis, Lacertidae)

Research on life-history traits of squamate reptiles has focused on North American species, while Asian taxa have been virtually ignored. In order to understand general patterns in reptile life histories, we need a broader data base. Our study on the slender-bodied lacertid lizard Takydromus septentrionalis provides the first detailed information on factors responsible for intraspecific variation in reproductive output and life history in a Chinese reptile. Clutches of recently collected lizards from five widely separated localities in China revealed major divergences in female body size at maturation, mean adult female body size, body condition after oviposition, size-adjusted fecundity, relative clutch mass, and mass and shape of eggs. Most of these geographical differences persisted when the same groups of females were maintained in identical conditions in captivity. Additionally, reproductive frequency during maintenance under laboratory conditions differed according to the animals' place of origin. Thus, the extensive geographical variation in reproductive and life-history traits that occurs within T. septentrionalis is exhibited even in long-term captives, suggesting that proximate factors that vary among localities (local conditions of weather and food supply) are less important determinants of life-history variation than are intrinsic (presumably genetic) influences. The maternal abdominal volume available to hold the clutch may be one such factor, based on low levels of variation in Relative Clutch Mass among populations, and geographical variation in the position of trade-off lines linking offspring size to fecundity.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 85 , 443–453.     

THE EVOLUTION OF BODY SHAPE IN RESPONSE TO HABITAT: IS REPRODUCTIVE OUTPUT REDUCED IN FLAT LIZARDS?

Body size and shape are primary determinants of reproductive output in a variety of taxa, so selection favoring specific body sizes and shapes may, in turn, have a direct affect on reproductive output, and ultimately fitness. In reptiles, species that occupy rocky habitats are often flattened, a morphological character that aids locomotion and life on rocks, but which may constrain reproductive output by reducing the amount of abdominal space available to fill with eggs or offspring. Using 20 species of tropical skink from a wide range of habitats, we quantified habitat use, body height, body volume, and reproductive output, to determine whether the evolution of a flattened body was correlated with a reduction in abdominal volume, and, in turn, with reduced reproductive output. In this group of lizards, the occupation of rocky habitats has led (1) to the evolution of a flattened body, and this shift in body shape has (2) caused a reduction in abdominal volume. Despite this reduction in abdominal volume reproductive output was unaffected, as flatter species compensate by being more "full" of eggs. Thus, we demonstrate that morphological adaptation for enhanced performance in specific habitats did not cause a reduction in instantaneous reproductive output.     

Maternal body-volume as a constraint on reproductive output in lizards: evidence from the evolution of viviparity

A few species of squamate reptiles contain both oviparous (egg-laying) and viviparous (live-bearing) populations, and thus offer exceptional opportunities to test adaptationist hypotheses on the determinants of reproductive output. We focus on the hypothesis that maternal body-volume constrains reproductive output in squamate reptiles. If females are “full” of eggs, what happens when viviparity evolves within a lineage? Eggs increase in volume and mass during development, primarily due to the uptake of water, so how can they be accommodated within the mother's abdomen? We predict that the resultant increase in relative clutch mass (RCM) will be lessened by (1) a decrease in reproductive output (by reducing the number or size of offspring), and/or (2) an increase in maternal body-volume (via modifications of size or shape of adult females). Our comparisons of conspecific oviparous and viviparous lizards (Lerista bougainvillii) confirm that live-bearers carry heavier clutches (in both absolute and relative terms) and show the predicted shifts in body size and shape of reproductive females. However, offspring size and number were unaffected by the evolution of viviparity, and the shifts in maternal morphology were too small to fully offset the increase in clutch mass. Thus, RCMs increased by 50%, indicating that viviparous females produced clutches which more completely filled the space available in the abdominal cavity. We conclude that maternal body-volume does play a role in determining reproductive output, but that the observed clutch masses may be optimized, rather than maximized, with respect to the abdominal space available.     

Maternal brooding in the children’s python <Emphasis Type="Italic">(Antaresia childreni)</Emphasis> promotes egg water balance

Parental care provides considerable benefits to offspring and is widespread among animals, yet it is relatively uncommon among squamate reptiles (e.g., lizards and snakes). However, all pythonine snakes show extended maternal egg brooding with some species being facultatively endothermic. While facultative endothermy provides thermal benefits, the presence of brooding in non-endothermic species suggests other potential benefits of brooding. In this study we experimentally tested the functional significance of maternal brooding relative to water balance in the children’s python, Antaresia childreni, a small species that does not exhibit facultative endothermy. Clutch evaporative water loss (EWL) was positively correlated with clutch mass and was much lower than expected values based on individual eggs. The conglomerate clutch behaved as a single unit with a decreasing surface area to volume ratio as clutch size increased. Maternal brooding had a dramatic impact on evaporation from eggs, reducing and possibly eliminating clutch EWL. In a separate experiment, we found that viability of unattended eggs is highly affected by humidity level, even in the narrow range from 75 to 100% relative humidity at 30.5°C (20–33 mg m⁻³ absolute humidity). However, the presence of the brooding female ameliorated this sensitivity, as viability of brooded clutches at 75% relative humidity was higher than that of non-brooded eggs at either the same absolute humidity or at near-saturated conditions. Overall, these results demonstrate that brooding behavior strongly promotes egg water balance (and thus egg viability) in children’s pythons.     

Ecological patterns of relative clutch mass in snakes

Summary Data on the relative clutch mass of snakes are summarized for over 100 populations. RCM was significantly lower in live bearing versus egg laying forms. We suggest that the longer reproductive season of viviparous snakes results in higher overall mortality compared to oviparous species; by reducing RCM, viviparous snakes may reduce this risk of mortality. Unlike lizards, no differences in RCM were found between categories of either escape behavior or foraging mode, possibly because detailed information on these behaviors are lacking for most snakes. In four populations examined, RCM did not vary among years. When compared to lizards, snakes demonstrate significantly higher RCM, perhaps owing to a more energetically efficient means of locomotion. Our data support the contention that RCM should be considered a separate and distinctive life-history characteristic of reptiles.     

Squamate phylogeny, taxon sampling, and data congruence

To investigate the affinities of snakes, amphisbaenians and dibamids, the phylogenetic relationships among the major lineages (families) of extinct and extant squamates are assessed through a combined analysis of 248 osteological, 133 soft anatomical, and 18 ecological traits. The osteological data set represents a revision of previous data, taking into account recent criticism; the ecological data set is new. In addition, potentially critical fossil taxa (polyglyphanodontids and macrocephalosaurs) are included for the first time. The osteological and soft anatomical data sets each place snakes within anguimorphs, with dibamids and amphisbaenians near gekkotans. The putative primitive fossil amphisbaenian Sineoamphisbaena groups with macrocephalosaurs and polyglyphanodontids, together the sister group to scleroglossans. All three data sets are congruent, and these results are reinforced by combined analyses. In these, as in the osteological analyses, snakes are nested within marine lizards. However, exclusion of fossil taxa from the osteological data set results in a ‘limbless clade’ consisting of snakes, amphisbaenians and dibamids, and introduces significant conflict between osteology and soft anatomy. Also, deletion tests and character weighting reveal that the signal in the reduced osteological data set is internally contradictory. These results increase confidence in the arrangement supported by the all-taxon osteological, the soft anatomical, and the combined data, and suggest that exclusion of fossils confounds the signal in the osteological data set. Finally, the morphological data support the nesting of snakes within marine lizards, and thus a marine origin of snakes. This result still holds when relationships between living forms are constrained to the topology suggested by molecular sequences: if marine lizards are allowed to ‘float’ within this molecular framework, they form the stem group to snakes, and do not group with varanids as previously suggested.See also Electronic Supplement at: http://www.senckenberg.de/odes/05-04.htm.     

Unraveling evolutionary lineages in the limbless fossorial skink genus Acontias (Sauria: Scincidae): are subspecies equivalent systematic units?

Subspecies in the limbless, endemic African fossorial skink genus Acontias constitute ill-defined operational taxonomic units, consequently considerable systematic debate has lingered on the systematic diversity within Acontias. In the present study, the systematic affinities among acontine taxa are explored with the utility of partial sequence data from two mitochondrial gene loci (16S rRNA and cytochrome oxidase subunit 1 (COI)) for all taxa, while two additional loci (12S rRNA, cytochrome b) were used to investigate relationships within the Acontias meleagris complex. Phylogenetic results, derived from the combined analysis, revealed two monophyletic clades. Clade 1 is comprised of small-bodied skinks while clade 2 comprised the medium bodied skinks. Within clade 2 none of the traditionally recognized subspecies formed reciprocally monophyletic groups. Furthermore, constraining the topology and enforcing sister taxa relationships between the assumed subspecies, consistently recovered a topology that was statistically significant worse, indicating that the traditionally designated subspecies groupings probably represent invalid taxonomic units, thus clearly reflecting considerable discord with current taxonomy. The burrowing life style of these lizards has probably led to marked convergent evolution and constrained the development of diagnostic morphological characters among these species. Morphological similarities in color as well as scale architecture within Acontias are labile and highly homoplaseous and do not reflect the evolutionary history of the group. Taxonomic implications of these results are discussed.     

WHY ARE CLUTCH SIZES MORE VARIABLE IN SOME SPECIES THAN IN OTHERS?

Animal species differ in the variability of their clutch sizes, as well as in mean clutch sizes. This phenomenon is particularly obvious in lizards, where virtually invariant clutch sizes have evolved independently in at least 23 lineages in seven families. Reduced variance in clutch size may arise either as an adaptation (because females with less variable clutch sizes have higher fitness) or as an indirect by-product of selection on other life-history characteristics. Comparative data on Australian scincid lizards indicate that variance in clutch sizes is lowest among species with low mean clutch sizes, small body sizes and a low variance in body sizes of adult females. Phylogenetic analysis shows that evolutionary decreases in the variance of clutch size have accompanied decreases in mean clutch sizes and decreases in the variance of adult female body sizes. Tropical lizards may also exhibit lower variance in clutch size. Most of these characteristics are correlated in occurrence, and may be allometrically tied to small body size. Hence, low variance in clutch size may be a consequence of allometric effects on a correlated suite of life-history characteristics. Exceptions to the general patterns noted above—especially, lizard species with invariant clutch sizes but large body sizes—may be due to loss of genetic variance for clutch sizes in lineages that have passed through a “bottleneck” of small body sizes and hence, low variance in clutch sizes.     

Morphological integration and adaptation in the snake feeding system: a comparative phylogenetic study

A long-standing hypothesis for the adaptive radiation of macrostomatan snakes is that their enlarged gape--compared to both lizards and basal snakes--enables them to consume "large" prey. At first glance, this hypothesis seems plausible, or even likely, given the wealth of studies showing a tight match between maximum consumed prey mass and head size in snakes. However, this hypothesis has never been tested within a comparative framework. We address this issue here by testing this hypothesis in 12 monophyletic clades of macrostomatan snakes using recently published phylogenies, published maximum consumed prey mass data and morphological measurements taken from a large sample of museum specimens. Our nonphylogenetically corrected analysis shows that head width--independent of body size--is significantly related to mean maximum consumed prey mass among these clades, and this relationship becomes even more significant when phylogeny is taken into account. Therefore, these data do support the hypothesis that head shape is adapted to prey size in snakes. Additionally, we calculated a phylogenetically corrected morphological variance-covariance matrix to examine the role of morphological integration during head shape evolution in snakes. This matrix shows that head width strongly covaries with both jaw length and out-lever length of the lower jaw. As a result, selection on head width will likely be associated with concomitant changes in jaw length and lower jaw out-lever length in snakes.     

Phylogeny of Philippine slender skinks (Scincidae: Brachymeles) reveals underestimated species diversity, complex biogeographical relationships, and cryptic patterns of lineage diversification

The spectacular, virtually endemic radiation of Philippine semi-fossorial skinks of the genus Brachymeles represent one of the few radiations of scincid lizards to possess both fully limbed and limbless species. And yet, nothing is known of the phylogenetic relationships of this exceptional group. Morphologically similar body plans have made it difficult to assess species-level diversity, and the genus has long been recognized as one of the more modest radiations of southeast Asian lizards. However, recent large-scale survey efforts have resulted in the discovery of numerous new species, and taxonomic studies indicate that the diversity within the genus Brachymeles is grossly underestimated. In this study we provide the first robust estimate of phylogenetic relationships within the genus Brachymeles using a multi-locus dataset and nearly complete taxonomic sampling. We provide statistical tests of monophyly for all polytypic species and two widespread limb-reduced species and our results indicate wholesale deviations from past summaries and taxonomic evaluations of the genus. With few exceptions, we are able to reject the monophyly of all polytypic and widespread species, thereby validating the need for large-scale taxonomic revisions. Our results reveal that the limbless, monotypic, genus Davewakeum is nested within Brachymeles. Mapping of body form on our preferred phylogenetic tree suggests that limb-reduction and digit loss has occurred on multiple occasions in the history of the genus. A Bayesian reconstruction of ancestral areas indicates strong statistical support for a minimum of five major dispersal events that have given rise to a major component of the observed species diversity on separate Pleistocene aggregate island platforms of the archipelago.     

Optimal offspring provisioning when egg size is "constrained": a case study with the painted turtle Chrysemys picta

Classic egg size theory predicts that, in a given environment, there is a level of maternal investment per offspring that will maximize maternal fitness. However, positive correlations among egg size and female body size are observed within populations in diverse animal taxa. A popular explanation for this phenomenon is that, in some populations, morphological constraints on egg size, such as ovipositor size (insects) or pelvic aperture width (lizards and turtles), limit egg size. Egg size may therefore increase with female body size due to body size‐specific constraints on investment per offspring, coupled with selection towards an optimal egg size. We use 17 years of data from a population of painted turtles Chrysemys picta to evaluate this hypothesis. In accordance with our predictions, we find that (1) morphological constraints on egg size are apparent only in relatively small females, similarly (2) egg mass exhibits a strong asymptotic relationship with female body size, suggesting egg mass is optimized only at large body sizes, (3) clutch size, not egg mass, varies with female condition, and (4) clutch size varies more than egg mass across years. Contrary to our predictions, we observe that (5) the egg mass‐clutch size tradeoff is not less pronounced at large body sizes. Our data do not fully support the traditional hypothesis, and recent models suggest that this hypothesis is indeed overly simplistic. When the selective environment of a female's offspring is influenced by her phenotype, optimal egg size may vary among maternal phenotypes. This concept can explain correlations among egg size and body size in many taxa, as well as the patterns observed in the present study. In this paradigm, a tight coupling of aperture width (or other ‘constraints’) and egg size may occur in small females, even when such morphological features are not causally related to variation in egg size. In this spirit, we question validity of invoking morphological constraints to explain covariation among egg size and female body size.     

Life-history evolution on tropidurinae lizards: influence of lineage, body size and climate

The study of life history variation is central to the evolutionary theory. In many ectothermic lineages, including lizards, life history traits are plastic and relate to several sources of variation including body size, which is both a factor and a life history trait likely to modulate reproductive parameters. Larger species within a lineage, for example tend to be more fecund and have larger clutch size, but clutch size may also be influenced by climate, independently of body size. Thus, the study of climatic effects on lizard fecundity is mandatory on the current scenario of global climatic change. We asked how body and clutch size have responded to climate through time in a group of tropical lizards, the Tropidurinae, and how these two variables relate to each other. We used both traditional and phylogenetic comparative methods. Body and clutch size are variable within Tropidurinae, and both traits are influenced by phylogenetic position. Across the lineage, species which evolved larger size produce more eggs and neither trait is influenced by temperature components. A climatic component of precipitation, however, relates to larger female body size, and therefore seems to exert an indirect relationship on clutch size. This effect of precipitation on body size is likely a correlate of primary production. A decrease in fecundity is expected for Tropidurinae species on continental landmasses, which are predicted to undergo a decrease in summer rainfall.