Overview

Brief Summary

Furcifer labordi is a sexually dimorphic chameleon species with an unusual annual life cycle. The entire population has a synchronized life history – all F. labordi individuals in a generation hatch, mature, and die in synchrony over the course of a single year. This species is unique among tetrapods because of its short post-hatching lifespan, lasting 4-5 months, and because it consequently spends more time inside of the egg than outside of it. Furcifer labordi inhabits coastal lowlands in southwest Madagascar, where it can be found on low perches (1-2 m above the ground) in bushes and trees. Common predators of this species include birds and snakes.

The social interactions of Furcifer labordi are physically aggressive, perhaps because of the high similarity in age and body size between individuals in a population. Several secondary sex characteristics are present. Most notably, males are distinguished by a high cranial casque and a prominent rostral process. Females are more colorful than males but their casques and rostral processes are highly reduced. The limited range and unique life cycle of this species may render it especially vulnerable to threats such as habitat destruction.

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Biology

This small reptile has one of the most unique life histories of all the four-limbed vertebrates (the tetrapods), living for just one year and spending eight to nine months of that time within an egg. As the first rains fall at the beginning of the wet season, around November, the Labord's chameleon eggs hatch. This synchronised hatching results in an entire population of Labord's chameleons that are roughly the same age. After hatching, Labord's chameleons have only four to five months of life left, the shortest known post-hatching lifespan of any tetrapod. They grow quickly, reaching sexual maturity in less than eight weeks, when they begin mating. Then, as swiftly as they have developed, Labord's chameleons begin to die-off, and by the time the dry season sets in, this amazing chameleon can no longer be seen; instead the entire species consists of eggs buried underground (6). This extreme life history is likely to be an adaptation to the extreme environment. In the arid and unpredictable region of Madagascar that this chameleon inhabits, it is a shrewd plan to stay within the relative stability and safety of an egg, underground, until conditions above ground are suitable (6). Chameleons are generally solitary, and move about on slender branches and twigs, which they grip with their fused toes. The prehensile tail provides an additional 'hand' on these precarious walkways as they scan the surrounding area with their independently mobile eyes for prey (4). Labord's chameleons normally sleep within two metres of the ground (6).
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Description

Known for its remarkable life history, Labord's chameleon is a fascinating and peculiar animal, with a compressed body, beautiful colouring, and turret-like eyes (4). The sexes differ significantly in appearance, with males being generally green with white stripes on the flanks, while females are considerably more colourful. The vivid green of the body is patterned with violet and blue markings on the flanks, and bright orange markings decorate the backbone. This contrasts with the striking red stripes on the skin of the throat. Male Labord's chameleons are also distinguished by their high, bony head crest (known as a casque), and the presence of a well-developed appendage jutting out from near the nostrils; this protuberance on females is only very small (2). The nasal projection and bony casque of the male enables females to recognise potential mates, and can be used as a weapon when in combat with other territorial males (4).
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Distribution

Continent: Indian-Ocean
Distribution: SE Madagascar  
Type locality: Cote occidentale de Madagascar
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Furcifer labordi is native to the western coast of Madagascar, in particular the dry forest habitats in the western and southwestern regions of the island. Its range extends from the lowland western forests from Soalala in the northwest to Ranobe in the southwest (Randrianantoandro et al. 2010).

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Range

This colourful chameleon is thought to be restricted to dry, western Madagascar (2).
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Physical Description

Morphology

Furcifer labordi is a medium-sized chameleon exhibiting substantial sexual dimorphism. In general, individuals of this species have relatively homogeneous scalation, disrupted only on the flanks where some larger scales are irregularly scattered. Prominent male secondary sex characteristics include a high cranial casque and a prominent, serrated dorsal crest consisting of enlarged conical scales. Canthi rostrales are present, and in males they merge into a prominent rostral appendage which is flat and fleshy rather than keratinized as in many other chameleon species. This rostral appendage is covered by relatively large, flat scales, and it may be under selection to become shorter and wider (see “Reproduction”). Less prominent gular and ventral crests are present in both sexes. In comparison to males, females have subtler or less developed dorsal crests, casques, and rostral appendages. Hemipenis morphology has not been formally documented in F. labordi, but the hemipenes of males of this species are thought to be ornamented with pedunculi and auriculae. Males are larger than females and have a swollen tail base. The karyotype of this species is 2n = 24 = (22 macrochromosomes and 2 microchromosomes; Grandidier 1872, Hechenbleikner 1942, Klaver and Böhme 1986, Necas 2004, Karsten et al. 2009c).

Males are green with a white dashed stripe along the medioventral line as well as white transverse stripes on the flanks. The body and tail are covered with vertical green and gray stripes. Females are much more colorful than males. Their green bodies are often dotted with brown, blue, and violet spots, and they may have a longitudinal row of spots on either flank that can be yellow or purple depending on the individual’s readiness to mate (see “Reproduction”). Females also possess orange vertebral markings and a violet-blue coloration on the flanks. The crosswise stripes on the flanks are also more visible in females than in males. Both sexes change color from green to brown or dark gray when stressed, but the lateral spots of females do not fade (Necas 2004, Karsten et al. 2009c).

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Size

Furcifer labordi is a medium-sized chameleon. In a population measured by Karsten et al. (2009b), adult male snout-vent lengths averaged 87.3 mm (n = 99, SE = 1.3) and adult female snout-vent lengths averaged 71.0 mm (n = 55, SE = 1.0). Necas (2004) reports that male total lengths (including the tail) may reach 308 mm, although females are generally smaller. Two females measured by Hechenbleikner (1942) had snout-vent lengths of 78 mm and 63 mm and tail lengths of 85 and 63 mm, respectively, for total lengths of 163 mm and 126 mm. Grandidier (1872) made the following measurements of the holotype. The length of the body is 111 mm, the length of the tail is 122 mm, and the length of the head, including the rostral process, is 36 mm. The rostral appendage is 12 mm long and 7 mm high. The casque has a maximum width of 13 mm and a maximum height of 32 mm (Grandidier 1872).

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Diagnostic Description

Distinctive anatomical features of F. labordi include its high casque and prominent, fleshy, and laterally flattened rostral appendage, both of which are much more strongly developed in males than in females. The coloration of this species (discussed in “Morphology” and “Reproduction”) may also help distinguish it from its closest relatives. In short, males are green with wide vertical gray stripes and narrow horizontal white stripes, and females are more colorful with their green background color decorated by vertical blue or violet stripes, rows of yellow or light purple spots, dots of brown, blue, and violet, and large spots of yellow, red, or violet near the head (Necas 2004, Karsten et al. 2009c).

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Look Alikes

This species is sometimes confused with the closely related species Furcifer antimena, F. rhinoceratus and F. angeli, although the height of F. labordi’s casque and the shapes and sizes of the rostral projections (tall and pointed straight ahead in F. labordi, as opposed to shorter and pointed upwards or downwards in the other species) may help distinguish this species. Furcifer labordi also closely resembles F. monoceras from the northeast region of Madagascar and Bradypodion xenorhinum from the Ruwenzori Mountains of Uganda and the Democratic Republic of the Congo. However, because F. labordi does not co-occur with many of these species in the wild, especially those from East Africa, confusion is only likely to arise with captive animals (Necas 2004).

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Ecology

Habitat

Habitat and Ecology

Habitat and Ecology

Laborde's Chameleon is strongly associated with remaining forest, especially spiny forest in the southwest and deciduous forest in the west (Raselimanana and Rakotomalala 2003). This species exhibits a bizarre annual life cycle where synchronous hatching occurs in November, followed by rapid growth into maturity, copulation in January and senescence and death by April (Karsten et al. 2008). Eggs are deposited in nests in the ground and remain in diapause until shortly before the rains (Karsten et al. 2008). There is little quantitative data on clutch size for this species in the wild, but a radio tracked female was observed laying 11 eggs in southern Madagascar (Karsten et al. 2008). This species is sexually dimorphic and exhibits physically intense combat and agnostic courtship (Karsten et al. 2008). In artificial combat scenarios, large males with short rostral appendages were the more successful whilst females appeared to show a preference for males with wide rostral appendages (Karsten et al. 2009). The species is prey for Madagascarophis and Mimophis snakes (Andriamandimbiarisoa 2007).


Systems
  • Terrestrial
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Furcifer labordi lives in warm coastal lowlands in western and southwestern Madagascar, where average temperatures are 22-28°C and the relative humidity is 80-100% year-round. This region is characterized by a long rainy season from November to March, which coincides with the active part of this species’ life cycle. The dry season is cooler and has little rainfall, a factor which may have influenced the evolution of the annual dormant phase in the life cycle of Furcifer labordi. This species prefers to roost in perches 1-2 m above the ground in dense thorny shrubs and small trees. Perch selection in F. labordi may be related to foraging opportunities and the avoidance of nocturnal predators (Necas 2004, Karsten et al. 2008, Randrianantoandro et al. 2010).

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Labord's chameleon inhabits small trees and thickets of thorny bushes (5).
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Associations

The available literature indicates that the main predators of chameleons are birds and snakes. Specifically, the Madagascar buzzard Buteo brachypterus and the colubrid snake Ithycyphus oursi have been documented preying on Furcifer chameleons, and the colubrid snakes Mimophis mahafaliensis and Madagascarophis meriodionalis are known to prey on F. labordi in particular (Jenkins et al. 2009).

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Population Biology

Because of its unusual annual life cycle, Furcifer labordi exhibits almost no age stratification within a population. All individuals in a population hatch, mature, and die together in a yearly cycle (Karsten et al. 2008).

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Life History and Behavior

Behavior

The social interactions of Furcifer labordi are characterized by intense physical combat, perhaps as a result of the similarity in body size among all individuals of a population due to its synchronized, annual life cycle. Furcifer labordi frequently prefaces both intra- and intersexual interactions with lateral displays, which may signal physical ability applicable to multiple social contexts. A lateral display involves perpendicular exposure of the laterally flattened body, where the tail is often coiled, to increase the apparent size of the chameleon. Interestingly, while many lizards bob their heads in territorial or threat displays, in F. labordi this behavior seems to be exclusively involved in courtship (Karsten et al. 2009c).

The best predictors of male fighting success are larger body size and shorter rostral appendages. During male-male combat, the primary mode of attack is biting. Thus, shorter rostral appendages, which interfere less with biting, may be beneficial for males during combat. However, intersexual selection may affect the rostral appendage differently by favoring increased width (see “Reproduction”). Male Furcifer labordi do not defend discrete territories and instead forage actively over relatively large home ranges. Radio-tracked individuals have been documented to cover up to 50 m in a single day (Karsten et al. 2009c).

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Cyclicity

The most unique aspect of this chameleon is its annual life cycle, in which all individuals in a population hatch, develop, reproduce, and die together in a period of 4-5 months. For the other 7-8 months of the year, populations of Furcifer labordi consist entirely of eggs (Karsten et al. 2008).

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Life Cycle

Furcifer labordi is unique among tetrapods for its annual life cycle. It spends the majority of its life inside the egg and has the shortest post-hatching lifespan of any known tetrapod. This life history resembles the life history of ephemeral insects or certain aquatic vertebrates more closely than it resembles that of other terrestrial tetrapods (Karsten et al. 2008).

During the dry season, populations of F. labordi consist entirely of eggs, which incubate for approximately 8 months before hatching simultaneously at the beginning of the rainy season in November. After hatching, F. labordi develops rapidly, reaching sexual maturity within 2 months and dying 2-3 months afterwards. Thus, this species spends more time inside the egg than out of it. The entire population grows synchronously, and no juveniles or adults are present when the eggs hatch (Karsten et al. 2008).

Reproduction occurs in January and February. Oviposition in this species has rarely been documented and is poorly understood, but observations suggest that female F. labordi dig underground burrows in which to lay their eggs. After reproduction, the adult population ages and dies, with the active season ending between February and April (Karsten and Andriamandimbiarisoa 2008, Karsten et al. 2008). See “Reproduction and Life History” for additional details.

Why Annualism?

There are currently two non-mutually exclusive hypotheses for why Furcifer labordi has developed its annual life cycle. One suggests that extreme life histories such as annualism may be adaptive in extreme environments. The region in Madagascar where this species occurs has a harsh and variable climate, with high variability in interannual rainfall. Life history theory suggests that such extreme seasonality is conducive to life history extremes such as highly “short-lived” or “long-lived” strategies. This hypothesis is supported by several mammals sympatric with F. labordi, whose life histories are more extreme than those of close relatives in environments with more moderate seasonality. Further supporting evidence comes from several lineages of fish such as killifish, in which phylogenetic analysis indicates that a short-lived, annual life cycle is probably ancestral but has been lost by species that live in more stable environments (Karsten et al. 2008).

A second hypothesis for the unusual life cycle of F. labordi is the relationship between high adult mortality and the evolution of rapid growth and earlier reproductive age. There tends to be a trade-off between resources allocated to maintenance of somatic cells, which contribute to longevity, and those allocated to reproduction. The physically intense social system of F. labordi, as well as the harsh environment in which it lives, drives evolutionary selection for reduced life span and smaller body size. Thus, rather than use valuable resources for extending longevity, this species may instead dedicate those resources to reproduction (Karsten et al. 2008).

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Life Expectancy

Eggs hatch approximately 8-9 months after they are laid. Sexual maturity is reached approximately 2 months after hatching, and the total post-hatching lifespan of this species is 4-5 months (Karsten et al. 2008).

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Reproduction

Reproduction in Furcifer labordi is oviparous and usually occurs within a short period of under 60 days. Females reproduce once in their year-long lifespan. Necas (2004) reports that clutches contain 8 eggs of about the same size as those of Furcifer lateralis. In what is currently the only published account of nesting behavior in wild F. labordi, Karsten and Andriamandimbiarisoa (2008) observed a female digging 138 mm into a sandy substrate, laying 11 eggs, and emerging from a second burrow, which she filled behind her while exiting. The total mass of the eggs was 4.4 g (over one-third of the female’s body mass before oviposition), and the average egg length was 11.7 mm (n = 11, SD = 0.37) (Karsten and Andriamandimbiarisoa 2008).

Courtship and Mate Selection

Females use body coloration as a signal of sexual receptiveness. Ordinarily, females of this species are emerald green with small, pale yellow spots on the lateral sides of the body. When females not receptive to mating are approached by a courting male, they darken their background color to dark green or black, making their yellow spots more conspicuous. Non-receptive females reject all male courtship, and male F. labordi generally spend little if any energy courting them. In contrast, in females that are potentially sexually receptive, the small yellow spots all become pastel violet, and a larger spot on the anterior flank turns red (Karsten et al. 2009c).

However, potentially receptive coloration does not guarantee a female’s willingness to mate. Some of these females remain passive and allow an approaching male to copulate with them, but others adopt an aggressive color pattern by darkening their background color to black with purple spots, a red flank spot, and orange spots along the dorsal crest. They may respond with physical aggression as well, which can include mouth gapes (opening the mouth widely to expose the color contrast of the interior mouth lining), lateral displays (presentation of the lateral flank with crests raised and lungs inflated to maximize apparent size), and gular displays (dorsoventral expansion of extendable skin below the jaw). They may also attack, lunge at, or even bite the male. All males observed by Karsten et al. (2009c) continued to court despite this, often nudging or stroking the female’s flank with their rostral appendages. While most of these males were still rejected, in some instances the female reverted to passive coloration and allowed copulation (Karsten et al. 2009c).

Potentially receptive Furcifer labordi females are more likely than females of the sympatric and closely related Furcifer verrucosus to respond aggressively to male courtship. However, males of F. labordi appear not to exhibit preferential mate selection but to be equally motivated to mate with all potentially receptive females. Due to this difference, it is likely that intersexual selection may affect male body size, secondary sex characteristics, and courting behavior. The best predictors of male mating success in F. labordi are body size and the width of the rostral appendage. Some have hypothesized that a wider rostral appendage may allow males to apply more lateral pressure to the female when courting, and if this pressure is sufficient it may induce her to switch from behavioral rejection to receptivity. However, longer rostral appendages are thought to interfere with biting, a central component of male-male combat. Thus, males of this species may experience intersexual and intrasexual selection for wider and shorter rostral appendages, respectively (Karsten et al. 2009c).

Additionally, while head-bob and pushup displays are common territorial signals in many male lizards, including F. verrucosus, in F. labordi these signals have only been observed during courtship. The extent to which the meanings of these signals might be malleable over evolutionary time in general remains unknown (Karsten et al. 2009c).

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Evolution and Systematics

Evolution

This species was originally described as Chamaeleo labordi (Grandidier 1872) and again as Chamaeleo barbouri (Hechenbleikner 1942). Klaver and Böhme placed it in the genus Furcifer in 1986, where it has remained since. Relationships within this genus, however, have not yet been decisively elucidated. There are currently no subspecies within Furcifer labordi (Necas 2004).

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Molecular Biology and Genetics

Genetics

Several nucleotide sequences from Furcifer labordi, including Mos, RAG-1, ND1, ND2, ND4, COI, several tRNAs, and partial sequences of the mitochondrial 12S and 16S rRNA genes, are available in the NCBI nucleotide database under accession numbers FJ984265.1, FJ984193.1, AF215138.1, AF443273.1, AF448767.1, and AF215264.1 (Vences 2000, Raxworthy et al. 2002, Townsend and Larson 2002, Townsend et al. 2009).

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Conservation

Conservation Status

IUCN Red List Assessment


Red List Category
VU
Vulnerable

Red List Criteria
B1ab(iii)

Version
3.1

Year Assessed
2011

Assessor/s
Jenkins, R.K.B., Andreone, F., Andriamazava, A., Anjeriniaina, M., Brady, L., Glaw, F., Griffiths, R.A., Rabibisoa, N., Rakotomalala, D., Randrianantoandro, J.C., Randrianiriana, J., Randrianizahana , H., Ratsoavina, F. & Robsomanitrandrasana, E.

Reviewer/s
Bowles, P. & Tolley, K.

Contributor/s

Justification
Listed as Vulnerable as the species has an extent of occurrence of 16,649 km in a region of southwestern Madagascar where its habitat continues to decline due to agricultural expansion and logging, and it is thought to occur as a severely fragmented population.

History
  • 1996
    Vulnerable (VU)
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The IUCN Red List of Threatened Species (1996) classified Furcifer labordi as Vulnerable.

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Status

Classified as Vulnerable on the IUCN Red List (1), and listed on Appendix II of CITES (3).
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Population

Population
In dry spiny forest habitats in south-western Madagascar this species occurs in population densities of 30.8 ha per ha (Karsten et al. 2009) whilst in the dry deciduous forests of the Menabe region in western Madagascar, it occurred at lower densities of 7.2 per ha (Randrianantoandro et al. 2010). In the southwest, the entire population of this species survives as eggs during the austral winter (Karsten et al. 2008) but it is not clear what life history is used by the populations further north, in less harsh environmental conditions. The patchy distribution of, and the severe human pressures placed on, remaining suitable habitat within this species' range suggest the population is both declining and severely fragmented.

Population Trend
Decreasing
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Threats

Major Threats

The forests in Madagascar's southwest are under high pressure from expanding rural and urban populations, and underwent some of the largest declines in forest cover between 1990 and 2000 in the whole of the island (Harper et al. 2007). The main threats to Laborde's Chameleon in this region are the conversion of native forest vegetation into charcoal and forest clearance for agriculture (Seddon et al. 2000, Andriamandimbiarisoa 2007). Similarly, the forests in western Madagascar are threatened by charcoal production and slash and burn clearance for subsistence agriculture, as well as timber harvesting (Smith et al. 1997, Ganzhorn et al. 2001, Young et al. 2008).There is no evidence that commercial collection currently poses a threat to this lizard.

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The primary threat to this species is the destruction, fragmentation, and degradation of the deciduous and spiny forests that it inhabits. Furcifer labordi may be particularly vulnerable because of its unusual annual life history: substantial mortality prior to reproduction could cause drastic population declines in a single generation. Karsten et al. (2009a) recommend protecting areas in the Ranobe region and north of the Fiherenana River in the Belalanda region to help conserve important populations of this species and the closely related and even rarer F. antimena (Karsten et al. 2009a).

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Like other chameleons of Madagascar, Labord's chameleon is likely to be facing two significant threats. Firstly, it may be impacted by exploitation for the wildlife trade, although this species is not one of the chameleon species that dominates Madagascar's chameleon trade (7). Secondly, it may be affected by the habitat degradation that is taking place over much of Madagascar. It is estimated that only 17 percent of the original vegetation of Madagascar remains, with agriculture, mining, and logging being among the main causes of habitat loss (8).
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Management

Conservation Actions

Conservation Actions

Laborde's chameleon occurs within the boundary of the Rserve Spciale d'Andranomena (Raselimanana and Rakotomalala 2003), Parc National Mikea and Parc National Kirindy Mitea (Raselimanana 2008) in the west, as well as in two other protected areas that are currently under development (Antimena in the Menabe Region and PK32 Ranobe in the Atsimo Andrefana Region). Additional survey work and research is needed into this species to better understand its current distribution, ecological requirements and life history. In particular, it is important to survey for this species in the forests north of the Tsiribihina River because, although comprehensive surveys have not foundLaborde's chameleon in Parc National Tsingy de Bemaraha (Randrianantoandro et al. 2008), it might occur in the lower altitude forest to the west. It is also important to ascertain whether the unusual life history documented for this lizard n the arid south is shared by subpopulations in the north, where environmental conditions are less harsh. Research is needed to establish the extent to which this species is threatened, and by which processes, in different parts of its range.

This species is listed on CITES Appendix II, but exports of wild caught individuals were suspended in 1994 (Carpenter et al. 2004). It is listed as a protected species (Category I, Class II) under Malagasy law which permits authorized collection from wild. Karsten et al. (2009) recommend that this species should be a conservation priority because of its restricted distributions, susceptibility to extirpation, low population densities, and lack of formal protection for habitat in the southwest.

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This species is listed in the CITES Appendix II.

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Conservation

Labord's chameleon is listed on Appendix II of the Convention on International Trade in Endangered Species (CITES) meaning that any international trade in this species should be carefully monitored (3). Otherwise, there are no specific conservation measures in place for this unique reptile, although numerous conservation organisations are working to conserve the remarkably diverse habitats of Madagascar and its intriguing animal inhabitants (9) (10) (11).
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Relevance to Humans and Ecosystems

Benefits

This species, like many chameleons, is popular among reptile hobbyists and breeders, although its post-hatching lifespan of only 4-5 months may make it less popular than longer-lived chameleons (Necas 2004). Additionally, this species has captured the interest of evolutionary biologists since the discovery of its unusual life cycle, which is unique among known tetrapods (Karsten et al. 2008).

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Wikipedia

Labord's chameleon

Labord's chameleon (Furcifer labordi) is a species of chameleon endemic to Madagascar. Like other Furcifer species (F. antimena, F. lateralis), it has an obligate year-long lifecycle. It lives for only about 4 to 5 months, making it the shortest lifespan ever recorded for a four legged vertebrate.[2][3] In their natural habitat, eggs hatch with the first rains in November, the growth is rapid, and adulthood is reached by January, at which time they breed. By later February or early March, females have deposited the eggs which will hatch next year, and the entire population dies until the next hatching.

In captivity, eggs of F. labordi have hatched after 4 months of incubation at 26 °C (79 °F). Juveniles grow very rapidly, reaching adulthood after 3 months. Females that were properly fed grew with eggs and a vivid coloration, whereas females that were fed a less caloric diet grew thinner and only showed a green coloration.

References[edit]

  1. ^ Andreone, F., Andriamazava, A., Anjeriniaina, M., Brady, L., Glaw, F., Griffiths, R.A., Jenkins, R.K.B., Rabibisoa, N., Rakotomalala, D., Randrianantoandro, J.C., Randrianiriana, J., Randrianizahana , H., Ratsoavina, F. & Robsomanitrandrasana, E. (2011). "Furcifer labordi". IUCN Red List of Threatened Species. Version 2012.1. International Union for Conservation of Nature. Retrieved 23 August 2012. 
  2. ^ Karsten, K. B.; Andriamandimbiarisoa, L. N.; Fox, S. F.; Raxworthy, C. J. (2008). "A unique life history among tetrapods: An annual chameleon living mostly as an egg" (PDF). Proceedings of the National Academy of Sciences 105 (26): 8980–8984. doi:10.1073/pnas.0802468105.  edit
  3. ^ Natural history: Nasty, brutish and short (subscription required)
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