Phenotypic responses of hatchlings to constant versus fluctuating incubation temperatures in the multi-banded krait, Bungarus multicintus (Elapidae)

Most studies on egg incubation in reptiles have relied on constant temperature incubation in the laboratory rather than on simulations of thermal regimes in natural nests. The thermal effects on embryos in constant-temperature studies often do not realistically reflect what occurs in nature. Recent studies have increasingly recognized the importance of simulating natural nest temperatures rather than applying constant-temperature regimes. We incubated Bungarus multicintus eggs under three constant and one fluctuating-temperature regimes to evaluate the effects of constant versus fluctuating incubation temperatures on hatching success and hatchling phenotypes. Hatching success did not differ among the four treatments, and incubation temperature did not affect the sexual phenotype of hatchlings. Incubation length decreased as incubation temperature increased, but eggs incubated at fluctuating temperatures did not differ from eggs incubated at constant temperatures with approximately the same mean in incubation length. Of the hatchling phenotypes examined, residual yolk, fat bodies and locomotor performance were more likely affected by incubation temperature. The maximal locomotor speed was fastest in the fluctuating-temperature and 30 degrees C treatments and slowest in the 24 degrees C treatment, with the 27 degrees C treatment in between. The maximal locomotor length was longest in the fluctuating-temperature treatment and shortest in the 24 degrees C and 27 degrees C treatments, with the 30 degrees C treatment in between. Our results show that fluctuating incubation temperatures do not influence hatching success and hatchling size and morphology any differently than constant temperatures with approximately the same mean, but have a positive effect on locomotor performance of hatchlings.     

Longterm effects of incubation temperatures on the morphology and locomotor performance of hatchling lizards (Bassiana duperreyi, Scincidae)

The phenotypes of hatchling reptiles are known to be affected by the thermal environments they experience during incubation, but the evolutionary and ecological significance of this phenotypic plasticity remains unclear. Crucial issues include: (i) the magnitude of effects elicited by thermal regimes in natural nests (as opposed to constant-temperature incubation); (ii) the persistence of these effects during ontogeny; and (iii) the consistency of these effects across different test conditions (does the thermal regime during embryogenesis simply shift the hatchling's thermal optimum for performance, or actually modify overall performance ability regardless of temperature?). We examined these questions by incubating eggs of scincid lizards (Bassiana duperreyi) from montane southeastern Australia, under two fluctuating-temperature regimes that simulated ‘cold’ and ‘hot’ natural nests. These thermal regimes substantially modified hatchling morphology (mass, body length, tail length, and the relationship between these variables), locomotor performance (running speeds over distances of 25 cm and lm), anti-predator ‘tactics’ and survival rates. The differences in locomotor performance persisted throughout the 20 weeks of our experiment. Lizards that emerged after ‘hot’ incubation were faster runners than their ‘cold’-incubated siblings under all thermal conditions that we tested. Thus, incubation temperatures modified overall locomotor ability, with only a minor effect on the set-point for optimum performance. The magnitude, persistence and consistency of these incubation-induced phenotypic modifications suggest that they may play an important role in evolutionary and ecological processes within lizard populations.     

Influence of incubation temperature on morphology, locomotor performance, and early growth of hatchling wall lizards (Podarcis muralis)

Eggs of wall lizards (Podarcis muralis) were incubated at three temperatures approaching the upper limit of viability for embryonic development in this species (26, 29, and 32 degrees C) to assess the influence of temperature on various aspects of hatchling phenotype likely affecting fitness. The thermal environment affected size and several morphometric characteristics of hatchling lizards. Hatchlings from eggs incubated at 32 degrees C were smaller (snout-vent length, SVL) than those from 26 and 29 degrees C and had smaller mass residuals (from the regression on SVL) as well as shorter tail, head, and femur relative to SVL. Variation in the level of fluctuating asymmetry in meristic and morphometric traits associated with incubation temperatures was quite high but not clearly consistent with the prediction that environmental stress associated with the highest incubation temperatures might produce the highest level of asymmetry. When tested for locomotor capacity in trials developed at body temperatures of 32 and 35 degrees C, hatchlings from the 32 degrees C incubation treatment exhibited the worst performance in any aspect considered (burst speed, maximal length, and number of stops in the complete run). Repeated measures ANCOVAs (with initial egg mass as covariate) of snout-vent length and mass of lizards at days 0 and 20 revealed significant effects of incubation temperature only for mass, being again the hatchlings from eggs incubated at 32 degrees C those exhibiting the smallest final size. All together, our results evidenced a pervasive effect of thermal regime during incubation (and hence of nest site selection) on hatchling phenotypes. However, incubation temperature does not affect hatchling phenotypes in a continuous way; for most of the analysed traits a critical threshold seems to exist between 29 and 32 degrees C, so that hatchlings incubated at 32 degrees C exhibited major detrimental effects. J. Exp. Zool. 286:422-433, 2000.     

Sex differences in optimal incubation temperatures in a scincid lizard species

Most theoretical models for the evolution of temperature-dependent sex determination (TSD) rely upon differential fitness of male and female offspring incubated under different thermal regimes. However, there are few convincing data on this topic. We studied incubation effects in a lizard species (Bassiana duperreyi, Scincidae) with genotypic sex determination, so that we could separate effects due to incubation temperatures from those due to offspring gender. We incubated eggs under two different fluctuating-temperature regimes that simulated hot and cold natural nest-sites. The effects of our incubation treatments on phenotypes of the hatchling lizards (morphology and locomotor performance) differed between the sexes. Females emerging from eggs exposed to the “hot nest” treatment (diel cycling, 23–31°C) were larger, and ran faster, than did their sisters from the “cold nest” treatment (16–24°C). Males showed a smaller and less consistent phenotypic response than females. These incubation-induced responses were relatively stable during the first few weeks of life post-hatching, at least in captive lizards maintained under laboratory conditions. These kinds of sex differences in the phenotypic responses of hatchling reptiles to incubation conditions provide a plausible basis for the evolution of temperature-dependent sex determination in reptiles. Received: 07 May 1998 / Accepted: 16 November 1998     

Influence of incubation temperature on hatchling phenotype in reptiles

Incubation temperature influences hatchling phenotypes such as sex, size, shape, color, behavior, and locomotor performance in many reptiles, and there is growing concern that global warming might adversely affect reptile populations by altering frequencies of hatchling phenotypes. Here I overview a recent theoretical model used to predict hatchling sex of reptiles with temperature-dependent sex determination. This model predicts that sex ratios will be fairly robust to moderate global warming as long as eggs experience substantial daily cyclic fluctuations in incubation temperatures so that embryos are exposed to temperatures that inhibit embryonic development for part of the day. I also review studies that examine the influence of incubation temperature on posthatch locomotion performance and growth because these are the traits that are likely to have the greatest effect on hatchling fitness. The majority of these studies used artificial constant-temperature incubation, but some have addressed fluctuating incubation temperature regimes. Although the number of studies is small, it appears that fluctuating temperatures may enhance hatchling locomotor performance. This finding should not be surprising, given that the majority of natural reptile nests are relatively shallow and therefore experience daily fluctuations in incubation temperature.     

Potential targets for selection during the evolution of viviparity in cold-climate reptiles

Viviparity (live-bearing) has evolved from oviparity (egg-laying) in more than 100 lineages of squamate reptiles (lizards and snakes). This transition generally has occurred in cool climates, where thermal differentials between eggs in the (cool) nest versus the (warm) maternal oviduct influence embryonic development, in ways that may enhance offspring fitness. To identify specific traits potentially under selection, we incubated eggs of a montane scincid lizard at conditions simulating natural nests, maternal body temperatures, and an intermediate stage (2-week uterine retention of eggs prior to laying). Incubation at maternal temperatures throughout incubation affected the hatchling lizard’s activity level and boldness, as well as its developmental rate, morphology, and locomotor ability. A treatment that mimicked the initial stages of the transition toward viviparity had a major effect on some hatchling traits (locomotor speeds), a minor effect on others (tail length, total incubation period) and no effect on yet others (offspring behaviors). More generally, different aspects of the phenotype are sensitive to incubation conditions at different stages of development; thus, the evolution of reptilian viviparity may have been driven by a succession of advantages that accrued at different stages of embryogenesis.     

Effects of constant and fluctuating temperatures on egg survival and hatchling traits in the northern grass lizard (Takydromus septentrionalis, Lacertidae)

To understand how nest temperatures influence phenotypic traits of reptilian hatchlings, the effects of fluctuating temperature on hatchling traits must be known. Most investigations, however, have only considered the effects of constant temperatures. We incubated eggs of Takydromus septentrionalis (Lacertidae) at constant (24 degrees C, 27 degrees C, 30 degrees C and 33 degrees C) and fluctuating temperatures to determine the effects of these thermal regimes on incubation duration, hatching success and hatchling traits (morphology and locomotor performance). Hatching success at 24 degrees C and 27 degrees C was higher, and hatchlings derived from these two temperatures were larger and performed better than their counterparts from 30 degrees C and 33 degrees C. Eggs incubated at fluctuating temperatures exhibited surprisingly high hatching success and also produced large and well-performed hatchlings in spite of the extremely wide range of temperatures (11.6-36.2 degrees C) they experienced. This means that exposure of eggs to adversely low or high temperatures for short periods does not increase embryonic mortality. The variance of fluctuating temperatures affected hatchling morphology and locomotor performance more evidently than did the mean of the temperatures in this case. The head size and sprint speed of the hatchlings increased with increasing variances of fluctuating temperatures. These results suggest that thermal variances significantly affect embryonic development and phenotypic traits of hatchling reptiles and are therefore ecologically meaningful.     

The influence of hydric environments during egg incubation on embryonic heart rates and offspring phenotypes in a scincid lizard (Lampropholis guichenoti)

Extensive evidence shows that incubation conditions can affect phenotypic traits of hatchling reptiles, but the relative importance of thermal versus hydric factors, and the proximate mechanisms by which such factors influence hatchling phenotypes, remain unclear for most species. We incubated eggs of an Australian scincid lizard, Lampropholis guichenoti, at four different moisture contents ranging from -500 to 0 kPa. Drier substrates reduced water uptake of eggs and resulted in smaller hatchlings, but other phenotypic traits (incubation periods, hatchling sex, body proportions, running speeds, growth rates post-hatching) were not affected by the hydric environment during incubation. Contrary to our prediction, lower water uptake during incubation (and hence, presumably, more viscous blood) did not affect embryonic heart rates. Thus, as in many other squamate species, hatchling phenotypes and embryonic developmental rates of L. guichenoti are less sensitive to hydric conditions in the nest than to thermal regimes.     

Phenotypic variation in an oviparous montane lizard (Bassiana duperreyi): the effects of thermal and hydric incubation environments

Recent studies have shown that incubation temperatures can profoundly affect the phenotypes of hatchling lizards, but the effects of hydric incubation environments remain controversial. We examined incubation-induced phenotypic variation in Bassiana duperreyi (Gray, 1938; Sauria: Scincidae), an oviparous montane lizard from south-eastern Australia. We incubated eggs from this species in four laboratory treatments, mimicking cool and moist, cool and dry, warm and moist, and warm and dry natural nest-sites, and assessed several morphological and behavioural traits of lizards after hatching. Incubation temperature influenced a lizard's hatching success, incubation period, tail length and antipredator behaviour, whereas variation in hydric conditions did not engender significant phenotypic variation for most traits. However, moisture affected incubation period slightly differently in males and females, and for a given snout-vent length moisture interacted weakly with temperature to affect lizard body mass. Although incubation conditions can substantially affect phenotypic variation among hatchling lizards, the absence of strong hydric effects suggests that hatchling lizards react less plastically to variation in moisture levels than they do to thermal conditions. Thus, our data do not support the generalization that water availability during embryogenesis is more important than temperature in determining the phenotypes of hatchling reptiles.     

Thermal regimes during incubation do not affect mean selected temperatures of hatchling lizards (Bassiana duperreyi, Scincidae)

Incubation temperatures profoundly affect many phenotypic traits of squamate reptiles, and mean selected body temperatures of such animals also are plastic in response to environmental factors. Plausibly, then, incubation temperatures might affect hatchling thermoregulation, either via adaptation (i.e., populations that historically experience different nest conditions, also will diverge in hatchling thermoregulatory behaviour) or phenotypic plasticity (incubation temperatures directly modify hatchling behaviours). We tested this hypothesis with a montane scincid lizard (Bassiana duperreyi), using thermal-imaging methods to quantify temperatures (of both head and body) selected by hatchling lizards. The young lizards kept their heads cooler than their bodies, but mean selected temperatures did not differ among hatchlings from three populations with differing thermal regimes in natural nests, nor were they affected by thermal conditions during incubation. The conservatism of mean selected temperatures stands in strong contrast to the lability of many other phenotypic traits in response to incubation temperatures in this species.