- Related Topics:
- chameleon
- mosasaur
- Iguania
- Autarchoglossa
- dammūsa
Life cycle
Most lizards reproduce by laying eggs. In some small species, the number of eggs is rather uniform for each laying or clutch. For example, all anoles (Anolis) lay but a single egg at a time, many geckos lay one or two eggs (depending upon the species), and some skinks have clutches of two eggs. A more general rule is that clutch size varies with the size, age, and condition of the mother. A clutch of four to eight eggs may be considered typical, but large lizards such as the iguanas may lay 50 or more eggs at one time. Lizard eggs are usually leathery-shelled and porous; they can expand by the absorption of moisture as the embryos grow. An exception occurs in the majority of egg-laying geckos, whose eggs have shells that harden soon after they are deposited and then show no further change in size or shape.
Embryonic development and sex determination
Viviparity, or the birthing of live young, occurs in some lizard species. For skinks, this is true for about one-third of the species, many of which live in tropical climates. In most other families that have live-bearing representatives, the species that are frequently exposed to cold conditions—either at high altitude or at extreme latitude—tend to be live-bearers. For example, all New Zealand geckos give birth to live young, yet all other geckos lay eggs. A great diversity of mechanisms exists that results in the production of live young. In some lizards the only difference between egg laying (oviparity) and live bearing (viviparity) is that shells never form around the “eggs.” The female retains them inside the oviduct until development is complete, and each egg already contains all of the energy necessary for development in its large yolk. In these cases, no additional nutrients pass from the mother to the offspring.
In other lizards, eggs released from the ovary contain most, but not all, of the energy necessary for development in the yolk. Several kinds of placentae can develop, depending on the species of lizard. The result is that some nutrients pass from the mother to the offspring during development. In just a few species, such as Mabuya heathi, tiny eggs with almost no yolk are released from the ovary and deposited in the oviduct. An advanced and complex placenta develops, and more than 99 percent of the nutrients required for embryonic development pass from the mother to the offspring. In these species the gestation period is usually very long ( 8–12 months).
Sex in most lizards is genetically and rigidly determined; a hatchling normally has either male or female reproductive structures. In representatives of most iguanian lizard families (Iguania) and in some species of whiptails, tegus, geckos, and skinks, the males have dissimilar sex chromosomes, comparable to the sex-chromosome system of most mammals. Some female geckos and wall lizards and all monitor lizards have sex-chromosome differences that are similar to those found in snakes. In a few lizard species (some iguanids, geckos, and wall lizards), no sex chromosomes exist. They rely on temperature-dependent sex determination (TSD)—that is, temperatures occurring within the nest during egg development control the sex of the hatchlings.
Parthenogenesis
Most lizard populations are evenly divided between females and males. Deviations from this pattern are found in parthenogenetic species, in which the young are produced from unfertilized eggs. Parthenogenesis in lizards was first discovered in all-female races of Lacerta in the Caucasus, but it is now known to occur in all-female species of whiptail lizards (Aspidoscelis) in the southwestern United States and parts of Mexico, several other Teiidae and Gymnophthalmidae (spectacled lizards or microteiids) in South America, and a few Gekkonidae. Parthenogenetic lizards appear to live in areas that are ecologically marginal for representatives of their genera. In Aspidoscelis and several other parthenogenetic species, convincing evidence exists that parthenogenetic forms arose through the hybridization of two bisexual species. The number of chromosomes in such species is usually double that in sexually reproducing species, but in a few cases, the number of chromosomes is triple. This results from the mating of a sexually reproducing species with one that is parthenogenetic. These offspring are called allotriploid because they represent a backcross that produces three sets of chromosomes.
Parental care
Parental care among lizards tends to be minimal following egg deposition, but there are striking exceptions. Many species dig holes in which the eggs are placed, whereas others bury them under leaf litter or deposit them in crannies of trees or caves. In contrast, females of some species, notably the five-lined skink (Eumeces fasciatus) of the United States and many of its relatives, remain with their eggs throughout the incubation time (about six weeks); they leave the clutch infrequently to feed. These skinks turn their eggs regularly and, if the eggs are experimentally scattered, will return them to the nest cavity. As soon as the young disperse, family ties are severed. Glass lizards (Ophisaurus, family Anguidae) appear to do the same thing. In addition, a number of viviparous lizards remove and eat the placental membranes from young when they are born.
In Australia, juvenile sleepy lizards (Tiliqua rugosa) remain in their mother’s home range for an extended period, and this behaviour suggests that they gain a survival advantage by doing so. Female sleepy lizards and those of the Baudin Island spiny-tailed skink (Egernia stokesii aethiops) recognize their own offspring on the basis of chemical signals. Consequently, parental care in lizards may be more widespread than previously thought. Nevertheless, since recognition systems are subtle, they are difficult to study.
Certain lizards, particularly some species of Gekkonidae, are known to be communal egg layers, with many females depositing their eggs at the same site. In addition, it appears that the same individual female may return to a particular site throughout her lifetime to deposit clutches of eggs. In Tropidurus semitaeniatus and T. hispidus, two species of South American ground lizards, females nest communally under slabs of rock situated on top of large boulders. In this specialized habitat, only a few appropriate nest sites are available, and thus they are limited resources. Males appear to take advantage of this situation, especially if nesting sites are located within their territories. It is likely that if a male defends a good nesting site, he should have access to more females than males who govern areas without high-quality nesting sites.
Maturation
Juvenile lizards are essentially miniature adults; they do not go through any larval phase or any other stage where they are dependent upon adults. They often differ from the adult in body colour or pattern and in certain body proportions. For example, the heads of hatchling lizards of some species tend to be proportionally larger than the heads of adults. Certain ornamental structures, such as the throat fan of the male green anole (Anolis) or the horns of some true chameleons (family Chamaeleonidae), develop as the lizards become sexually mature. The tails of juveniles in many lizard species are coloured differently from those of adults. Juvenile tails are brilliant blue, orange, or red and easily discarded (autotomized) when escaping a predator. Tail colour usually changes when the lizards reach sexual maturity.
Some of the smaller lizards mature very quickly, and population turnover (that is, the replacement of one generation by another) is essentially an annual event. For example, in the small, side-blotched lizard (Uta stansburiana) of western North America, the young hatch in July and reach sexual maturity the following autumn. At this time, males undergo spermatogenesis and mating takes place. Female side-blotched lizards accumulate large quantities of fat, which appear to be utilized in the production of eggs the following spring. Adult mortality in this species is 90 percent or more per year and may be a result of predation, inclement weather, or other factors. Conversely, the population dynamics of a single species living under a variety of environmental conditions may be very different from one region to another. For example, in areas with long winters where lizards experience long periods of hibernation, they may have greater longevity and slower population turnover.
On the other hand, large lizards may take several years to reach sexual maturity, and little information exists on the dynamics of natural populations of most lizard species. In captivity, many species are long-lived. Gila monsters (Heloderma) have been kept in captivity for more than 25 years, and even some small geckos have been kept for as long as 20 years. There is a report of a 46-year-old captive male slowworm (Anguis fragilis) mating with a 20-year-old female.
Ecology
Thermoregulation
The most important environmental variable to a lizard is almost certainly temperature. Like fish and amphibians, lizards are ectothermic; they receive heat from their surroundings. Although the term cold-blooded is typically applied to such organisms, it is a misnomer. The blood of lizards is not cold unless the lizard is cold. Under conditions where normal activities occur, lizard blood is as warm as or warmer than that of mammals. Nevertheless, all temperatures are not equally acceptable to lizards. Most species seek out relatively specific body temperatures, called “preferred temperatures,” that mostly range from 28 to 38 °C (82 to 100 °F).
Although metabolic energy is not utilized to control body temperature, considerable thermoregulation is accomplished through behavioral means, if the lizard has a choice. Typically, a diurnal lizard emerges early in the morning and suns itself, orienting the body to maximize exposure to the sun, until the preferred temperature is achieved. The ability to absorb heat from solar radiation may permit the lizard to warm itself well above air temperatures. For example, Liolaemus multiformis, a small lizard that lives high in the Andes, has the ability to raise its body temperature to 35 °C (95 °F) while air temperatures are at 10 °C (50 °F) or lower.
The preferred body temperature plays a critical physiological role in the life of a lizard. All physiological processes are temperature-dependent, and physiological function influences behaviour. In most instances, the lizard’s “performance,” (that is, the lizard’s ability to execute various behaviours or function well metabolically) is optimal within a small range of temperatures. To maximize performance, the lizard should seek to maintain its body temperature within this temperature range when at all possible.
Traditionally, the immediate environment in which a lizard lives has been considered the primary determinant of the lizard’s body temperature; however, since thermoregulation is complex, there are constraints. Lizards living in hot deserts might be expected to be active at higher body temperatures than those living in well-shaded tropical habitats. Nonetheless, a combination of factors including evolutionary history, the immediate thermal conditions, and the “costs” associated with behavioral thermoregulation determines temperatures at which a lizard will operate.
The effect of evolutionary history is obvious when comparing certain groups of lizards. All whiptail lizards and racerunners in the genera Aspidoscelis and Cnemidophorus are active at body temperatures between 37 and 40 °C (99 and 104 °F) whether they live in the hottest part of the Mojave Desert of southern California or along trails in the Amazon Rainforest. In addition, all lizards in the family Xantusiidae, a group distributed from the Mojave Desert southward through the rainforests of Central America, are active at body temperatures between 25–28 °C (77–82 °F). Whiptails adjust their activity periods to take advantage of heat sources in environments where temperatures are relatively low, whereas the tiny desert night lizard (Xantusia vigilis) occupies a microhabitat that remains cool in an otherwise hot place. Although some desert lizards have slightly higher body temperatures than their close relatives in more moderate habitats, the immediate thermal conditions often determine when and where a lizard will be active rather than what its body temperature will be.
Several costs to thermoregulation exist, but only a few have been studied. Time spent basking to gain heat or escaping extremely high or extremely low temperatures cannot be used for feeding or reproduction. Basking in direct sunlight to gain heat places a lizard in an exposed location where predators can capture it. Lizards whose body temperatures are outside of the optimal range for their species may not perform as well in social interactions as those lizards at optimal body temperatures. Some of the lesser-known costs include reduced growth rates and longer time to sexual maturity, increased incubation times for eggs or embryos when optimal temperatures cannot be reached, and a reduced ability to escape falling temperatures, which may result in the freezing of body tissues.
