Viviparous lizard | |
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Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Reptilia |
Order: | Squamata |
Family: | Lacertidae |
Genus: | Zootoca |
Species: | Z. vivipara |
Binomial name | |
Zootoca vivipara (Lichtenstein, 1823) [2] | |
The viviparous lizard is found across Northern Europe through Central Asia, ranging further north than any other land-dwelling reptile. | |
Synonyms | |
Lacerta viviparaLichtenstein, 1823 |
The viviparous lizard, or common lizard, (Zootoca vivipara, formerly Lacerta vivipara) is a Eurasian lizard. It lives farther north than any other species of non-marine reptile, and is named for the fact that it is viviparous, meaning it gives birth to live young (although they will sometimes lay eggs normally). [3] Both "Zootoca" and "vivipara" mean "live birth", in (Latinized) Greek and Latin respectively. It was called Lacerta vivipara until the genus Lacerta was split into nine genera in 2007 by Arnold, Arribas & Carranza. [4]
Male and female Zootoca vivipara are equally likely to contract blood parasites. [5] Additionally, larger males have been shown to reproduce more times in a given reproductive season than smaller ones. [6]
The lizard is also unique as it is exclusively carnivorous, eating only flies, spiders, and insects. [7] Studies show that the more carnivorous an individual is (the more insects they eat), the less diverse the population of parasitic helminths that infest the lizards. [7]
Zootoca vivipara lives in very cold climates, yet participates in normal thermoregulation instead of thermoconformity. [8] They have the largest range of all terrestrial lizards which even include subarctic regions. It is able to survive these harsh climates as individuals will freeze in especially cold seasons and thaw two months later. They also live closer to geological phenomena that provide a warmer environment for them. [8]
Zootoca vivipara is a small lizard, with an average length between {150-200 mm} . [3] They exhibit no particular colour, but can be brown, red, grey, green, or black. [3] The species exhibits some sexual dimorphisms. Female Z. vivipara undergo colour polymorphism more commonly than males. A female lizard's display differs in ventral coloration, ranging from pale yellow to bright orange and a mixed coloration. There have been many hypotheses for the genetic cause of this polymorphic coloration. These hypothesis test for coloration due to thermoregulation, predator avoidance, and social cues, specifically sexual reproduction. Through an experiment conducted by Vercken et al., colour polymorphism in viviparous lizard is caused by social cues, rather than the other hypotheses. More specifically, the ventral coloration that is seen in female lizards is associated with patterns of sexual reproduction and sex allocation. [9]
The underside of the male is typically more colourful and bright, with yellow, orange, green, and blue, and the male typically has spots along its back. [3] On the other hand, females typically have darker stripes down their backs and sides.[ citation needed ] Additionally, males have been found to have larger heads than their female counterparts, and this trait appears to be sexually selected for. [6] Males with larger heads are more likely to be successful in mating and male-male interactions than smaller-headed Z. vivipara. [6] Larger males also have been shown to reproduce more frequently during one mating season compared to smaller males. [6] Characteristic behaviors of the species includes tongue flicking in the presence of a predator and female-female aggression that seems to be mediated by the colour of their side stripe. [3]
Z. vivipara is terrestrial, so they spend most of their time on the ground, though they do occasionally visit sites of higher elevation. [10] The lizard thermoregulates by basking in the sun for much of the time. In colder weather, they have been known to hibernate to maintain proper body temperatures. They hibernate between October and March. [11] Their typical habitats include heathland, moorland, woodland and grassland. [12]
The viviparous lizard is native to much of northern Eurasia. In Europe, it is mainly found north of the Alps and the Carpathians, including the British Isles but not Iceland, as well as in parts of northern Iberia and the Balkans; In Asia it is mostly found in Russia, excluding northern Siberia, and in northern Kazakhstan, Mongolia, China, and Japan.[ citation needed ] Z. vivipara has the largest distribution of any species of lizard in the world. [13]
The size of the home range of the lizard ranges from 539 m2 to 1692 m2, with males generally having larger home ranges. [3] The size of an individual lizard's home range is also dependent on population density and the presence of prey. [3]
Unlike many other lizards, Z. vivipara is exclusively carnivorous. [7] Their diet consists of flies, spiders, and various other insects, including hemipterans (such as cicadas), moth larvae, and mealworms. [7] [14] The species is a predator, so it actively hunts down all of its prey. [7] One study found that when controlled for body size, females consumed more food than males. Feeding rates also increased with increased sunshine. [15]
Birds are common predators of Z. vivipara. Male-biased predation of Z. vivipara by the great grey shrike (L. excubitor) has been studied, finding that adult males, over adult females and juveniles, were preferentially predated on. This bias may be due to increased activity of adult males during the reproductive season. [16]
Predators of this species include birds of prey, crows, snakes, shrikes, hedgehogs, shrews, foxes, and domestic cats. [17] [11]
Z. vivpara can be infested by helminths, a small parasitic worm. [7] The species diversity of parasites is affected by the diet of the individual lizard and the number of parasites on a host is affected by the host's size. [7] Results of a study shows that the more carnivorous an individual is, the less diverse its parasite population. Additionally, larger lizards had a greater number of parasites on them. [7]
Z. vivipara is also infected by blood parasites. In a study investigating the prevalence of blood parasites in Z. vivipara and L. agilis, Z. vivipara was found to be parasitized with an incidence rate of 39.8%, while L. agilis was parasitized with an incidence rate of 22.3%. [5] This same study shows that there was not a significant difference between the parasitization of male and female Z. vivipara. [5]
The viviparous lizard is named as such because it is viviparous. This refers to its ability to give birth to live young, although the lizards are also able to lay eggs. [18] The origin of this characteristic is under debate. Some scientists argue that viviparity evolved from oviparity, or the laying of eggs, only once. [18] Proponents of this theory also argue that if this is the case, it is possible, though rare, for species to transition back to oviparity. [18] Research from Yann Surget-Groba suggests that there have in fact been multiple events of the evolution of viviparity from oviparity across different clades of the viviparous lizard. They also argue that a reversion to oviparity is not as rare as once believed, but has occurred 2 to 3 times in the history of the species. [18]
The range of viviparous populations of Z. vivipara extends from France to Russia. Oviparous populations are only found in northern Spain and the southwest of France. [19] Some research in the Italian alps has suggested that distinct populations of oviparous and viviparous Z. vivipara should be considered separate species. Cornetti et al. (2015) identified that viviparous and oviparous subpopulations in contact with each other in the Italian alps are reproductively isolated. [20] Hybridization between viviparous and oviparous individuals of Z. vivipara leads to embryonic malformations in the laboratory. [21] However, these crosses do produce a "hybridized" generation of offspring, with females retaining embryos for much longer in utero than oviparous females, with embryos surrounded by thin, translucent shells. [22] [23]
Z. vivipara juveniles reach sexual maturity during their second year of their life. [24] A study that explored the presence of male sex cells in reproducing males found that for the two weeks following the end of hibernation, males are infertile, and therefore incapable of reproducing. [24] The same study also found that larger males produce more sperm during the reproductive season and have fewer left over at the end of the reproductive season than their smaller counterparts. [24] This suggests that the larger a male is, the more reproductive events they participate in. [24]
Research also suggests that in exclusively oviparous populations of Z. vivipara, altitude influences the number of clutches laid in a reproductive season as well as when reproduction begins. Generally, lizards living at higher altitudes have been found to begin reproduction later and lay fewer clutches (often 1) in a given reproductive season. [24]
Z. vivipara typically lives for 5 to 6 years.[ citation needed ]
Head size is a sexually dimorphic trait, with males having larger heads than females. The average head width and length of the males measured were found to be 5.6 and 10.5 mm (0.22 and 0.41 in), respectively. [6] The average head width and length of the females measured were found to be 5.3 and 9.7 mm (0.21 and 0.38 in), respectively. During the first state of courtship in Z. vivipara, called "Capture", the male uses its mouth and jaw to capture the female and initiate copulation. [6] The results of this study demonstrated that males with larger head sizes (both length and width) were more successful in mating than those with smaller heads, suggesting that head size undergoes sexual selection. [6]
Head size has also been shown to be a predictor of success in male-male interactions. [6] The head is used as a weapon in male-male interactions, and a larger head is typically more effective, leading to greater success during male-male aggressive encounters. [6] This aggression and interaction is centered around available mates, so males with smaller heads have significantly less access to females for reproduction. [6]
This lizard has an exceptionally large range that includes subarctic geography. [8] As a result, thermoregulation is necessary for the thermal homeostasis of the species. Typically, in temperature extremes, a species will adopt the behavioral strategy of thermoconformity, [8] where they do not actively thermoregulate, but adapt to survive in the harsh temperature. This occurs because the cost of thermoregulating in such an extreme environment becomes too high and begins to outweigh the benefits. [8] Despite this, Z. vivipara still employs the strategy of thermoregulation, like basking. [8] Thermoregulation is important in Z. vivipara as it allows for proper locomotive performance, escape behavior, and other key behaviors for survival. [8] The ability of Z. vivipara to thermoregulate in such harsh environments has been attributed to two primary reasons. [25] The first is that Z. vivipara has remarkable behaviors to combat the cold, and there are geological phenomena in their distribution that maintains their habitats at a temperature that the species can survive in. [25] One of the specific behaviors used to combat the extreme cold is a "supercooled" state. [25] Z. vivipara remains in this state through the winter until temperatures dropped below −3 °C (27 °F). After that, individuals completely froze until they were thawed by warmer weather later in the year, often 2 months later. [25] Despite very cold air in the subarctic habitats of these lizards, the soil-heating effects of unfrozen groundwater has been observed regulating the temperature of their soil habitats. They find warm microhabitats that do not drop below the freezing point of their body fluids. These lizards have exceptional hardiness to the cold, which allows them to hibernate in upper soil layers in temperatures as low as −10 °C (14 °F). This cold hardiness along with the favorable hydrogeological conditions of groundwater-warmed soil habitats allows for the wide distribution of lizards throughout the palearctic. [26]
The colour polymorphism of female Z. vivipara has not been thoroughly studied in past years, regardless of the extensive research done on the species itself. [27] Females exhibit three types of body colouration within a population: yellow, orange, and mixture of the two. These discrete traits are inherited maternally and exist throughout the individual's lifetime. [27] The organism's colour morphs are determined by their genotype as well as their environment.
The frequency of multiple morphs occurring in a population varies with the level of population density and frequency-dependent environments. [27] These factors cause the lizards to vary in terms of their fitness (clutch size, sex ratio, hatching success). [27] In lower density populations, colour polymorphism is more prevalent. [9] This is because viviparous lizards thrive in environments where intraspecific competition is low. [28] Increased competition among individuals results in lower survival rates of lizards. Additionally, female lizards disperse through habitats based on the frequency of colour types that are already present in the population. [28] Their reproductive abilities vary according to this frequency-dependent environment. The number of offspring that they produce correlates with the colour morph: yellow females produce the fewest offspring, while orange females produce more than yellow, but fewer than mixed females, which produce the most offspring. [9] The amount of offspring produced varies in regards to colour frequencies in the population; for example, if yellow females have higher density within the population, the clutch size for orange lizards is usually lower. [9]
Orange females are more sensitive to intraspecific and colour-specific competition. [9] They have smaller clutch sizes when the density of the population is high, or when the number of yellow females in the population is high. This could be due to their need to conserve energy for survival and reproductive events. [9] Their colour morph remains in the population due to the trade-off between the size of offspring and the clutch size. Offspring born in smaller clutches are often larger and thus have a higher survival likelihood. [9] Natural selection will favor individuals with larger size because of their advantage in physical competition with others. Yellow females have larger clutch sizes early in their life, but their hatch success decreases as the female ages. [9] Their reproductive viability decreases, resulting in fewer offspring throughout their lifetime. Yellow morphs remain in the population due to their large clutch size, which causes an increased frequency of those females. [9] Selection favors the yellow morph because of the ability to produce large clutch sizes, which increases the female's fitness. In mixed-coloured females, reproductive success is less sensitive to competition and frequency-dependent environments. [9] Since these lizards show a mixture of yellow and orange colouration, they adopt benefits from both of the morphs. As a result, they can maintain high reproductive success and hatching success with large clutch sizes. [9] Their colour morph remains in the population due to its high fitness, which selection will favor.
All three colours have evolutionary advantages in different ways. While yellow females have higher fitness due to their large clutch sizes, orange females enjoy high fitness due to their large body size and increased competitive advantages. Mixed females exhibit both of these advantages.
Gestation is the period of development during the carrying of an embryo, and later fetus, inside viviparous animals. It is typical for mammals, but also occurs for some non-mammals. Mammals during pregnancy can have one or more gestations at the same time, for example in a multiple birth.
Birth is the act or process of bearing or bringing forth offspring, also referred to in technical contexts as parturition. In mammals, the process is initiated by hormones which cause the muscular walls of the uterus to contract, expelling the fetus at a developmental stage when it is ready to feed and breathe.
Lacerta is a genus of lizards of the family Lacertidae.
In animals, viviparity is development of the embryo inside the body of the mother, with the maternal circulation providing for the metabolic needs of the embryo's development, until the mother gives birth to a fully or partially developed juvenile that is at least metabolically independent. This is opposed to oviparity, where the embryos develop independently outside the mother in eggs until they are developed enough to break out as hatchlings; and ovoviviparity, where the embryos are developed in eggs that remain carried inside the mother's body until the hatchlings emerge from the mother as juveniles, similar to a live birth.
Ovoviviparity, ovovivipary, ovivipary, or aplacental viviparity is a term used as a "bridging" form of reproduction between egg-laying oviparous and live-bearing viviparous reproduction. Ovoviviparous animals possess embryos that develop inside eggs that remain in the mother's body until they are ready to hatch.
An egg is an organic vessel grown by an animal to carry a possibly fertilized egg cell and to incubate from it an embryo within the egg until the embryo has become an animal fetus that can survive on its own, at which point the animal hatches.
The sand lizard is a lacertid lizard. There are several subspecies, including L. a. agilis, L. a. argus, and L. a. exigua.
Podarcis muralis is a species of lizard with a large distribution in Europe and well-established introduced populations in North America, where it is also called the European wall lizard. It can grow to about 20 cm (7.9 in) in total length. The animal has shown variation in the places it has been introduced to. Fossils have been found in a cave in Greece dating to the early part of the Holocene.
Fish reproductive organs include testes and ovaries. In most species, gonads are paired organs of similar size, which can be partially or totally fused. There may also be a range of secondary organs that increase reproductive fitness. The genital papilla is a small, fleshy tube behind the anus in some fishes, from which the sperm or eggs are released; the sex of a fish can often be determined by the shape of its papilla.
Oviparous animals are animals that reproduce by depositing fertilized zygotes outside the body in metabolically independent incubation organs known as eggs, which nurture the embryo into moving offsprings known as hatchlings with little or no embryonic development within the mother. This is the reproductive method used by most animal species, as opposed to viviparous animals that develop the embryos internally and metabolically dependent on the maternal circulation, until the mother gives birth to live juveniles.
The common side-blotched lizard is a species of side-blotched lizard in the family Phrynosomatidae. The species is native to dry regions of the western United States and northern Mexico. It is notable for having a unique form of polymorphism wherein each of the three different male morphs utilizes a different strategy in acquiring mates. The three morphs compete against each other following a pattern of rock paper scissors, where one morph has advantages over another but is outcompeted by the third.
Iberolacerta cyreni, commonly known as the Cyren's rock lizard, is a species of lizard in the family Lacertidae. The species is endemic to central Spain and is currently listed as endangered by the IUCN due to global warming. I. cyreni has evolved to exhibit key behavioral characteristics, namely individual recognition, in which a lizard is able to identify another organism of the same species, as well as thermoregulation.
The eastern three-lined skink, also known commonly as the bold-striped cool-skink, is a species of skink, a lizard in the family Scincidae. The species is endemic to Australia. A. duperreyi has been extensively studied in the context of understanding the evolution of learning, viviparity in lizards, and temperature- and genetic-sex determination. A. duperreyi is classified as a species of "Least Concern" by the IUCN.
Saiphos equalis, commonly known as the yellow-bellied three-toed skink or simply three-toed skink, is a species of burrowing skink found in eastern Australia. It is the only species classified under the genus Saiphos.
Sexual selection in scaled reptiles studies how sexual selection manifests in snakes and lizards, which constitute the order Squamata of reptiles. Each of the over three thousand snakes use different tactics in acquiring mates. Ritual combat between males for the females they want to mate with includes topping, a behavior exhibited by most viperids in which one male will twist around the vertically elevated fore body of its opponent and forcing it downward. It is common for neck biting to occur while the snakes are entwined.
Ctenophorus decresii, also known commonly as the tawny dragon or the tawny crevice-dragon, is a species of lizard in the family Agamidae. The species is endemic to Australia. The average snout-to-vent length (SVL) of the species is 80.76 mm (3.180 in) with larger individuals being around 89 mm (3.5 in) and smaller individuals around 72 mm (2.8 in). The optimal time for mating in this species is two to three weeks after the females emerge from hibernation. Eggs are typically laid from September to October with most of them being laid earlier in the period. C. decresii is known for its variations in throat colours which change based on environmental conditions. Its primary food sources consist of both vegetation and invertebrates, and it prefers to live in rocky habitats.
Animals make use of a variety of modes of reproduction to produce their young. Traditionally this variety was classified into three modes, oviparity, viviparity, and ovoviviparity.
Vertebrate maternal behavior is a form of parental care that is specifically given to young animals by their mother in order to ensure the survival of the young. Parental care is a form of altruism, which means that the behaviors involved often require a sacrifice that could put their own survival at risk. This encompasses behaviors that aid in the evolutionary success of the offspring and parental investment, which is a measure of expenditure exerted by the parent in an attempt to provide evolutionary benefits to the offspring. Therefore, it is a measure of the benefits versus costs of engaging in the parental behaviors. Behaviors commonly exhibited by the maternal parent include feeding, either by lactating or gathering food, grooming young, and keeping the young warm. Another important aspect of parental care is whether the care is provided to the offspring by each parent in a relatively equal manner, or whether it is provided predominantly or entirely by one parent. There are several species that exhibit biparental care, where behaviors and/or investment in the offspring is divided equally amongst the parents. This parenting strategy is common in birds. However, even in species who exhibit biparental care, the maternal role is essential since the females are responsible for the incubation and/or delivery of the young.
The temperature-size rule denotes the plastic response of organismal body size to environmental temperature variation. Organisms exhibiting a plastic response are capable of allowing their body size to fluctuate with environmental temperature. First coined by David Atkinson in 1996, it is considered to be a unique case of Bergmann's rule that has been observed in plants, animals, birds, and a wide variety of ectotherms. Although exceptions to the temperature-size rule exist, recognition of this widespread "rule" has amassed efforts to understand the physiological mechanisms underlying growth and body size variation in differing environmental temperatures.
Bimodal reproduction is the ability of a vertebrate animal to reproduce by both laying eggs and giving birth. Examples of these animals include Saiphos equalis and Zootoca vivipara,