Liolaemus chiliensis

Summary

Liolaemus chiliensis (Chilean tree iguana) is a species of lizard in the family Liolaemidae, also referred to as the weeping or crying lizard in English. Synonyms for this species include Liodeira chilensis and Calotes chiliensis.[1] Less commonly, it is called the Talcahuano Smooth-throated Lizard.[2]

Liolaemus chiliensis
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Order: Squamata
Suborder: Iguania
Family: Liolaemidae
Genus: Liolaemus
Species:
L. chiliensis
Binomial name
Liolaemus chiliensis
(Lesson, 1830)

This species is well known for its distress calls, although the sound of these calls does not resemble weeping or crying.

The specific name chiliensis is derived from the species' country of origin, Chile. It is endemic to central Chile and southwestern Neuquen Province, Argentina. This iguanid lizard is found inhabiting bushes and branches in open forests and ecotonal scrublands or scrub-steppes, chiefly in the Chilean Matorral ecoregion.[3]

Taxonomy edit

Many previous phylogenetic studies for L. chiliensis have been incomplete; a more recent biogeographic analysis from 2006 reconstructs the phylogenetic tree of the lizard's genus and resolves controversies about subgrouping of species or subspecies within L. chiliensis.[4] The authors of this study group L. chiliensis with L. curicensis, L. nitidus, L. robertmensi, L. saxatillis, L. gracilis, and L. tandilensis under the robertmensi group.[4] Sources differ in whether they refer to Liolaemus chiliensis as a single species or group of species that includes the majority of species in the genus Liolaemus.[5] Some have distinguished multiple species within L. chiliensis due to differences in "snout-vent length, squamation, and dorsal and ventral colors."[6] Much of the debate revolves around the difference between a subspecies and full species. The Liolaemus genus includes over 160 species, and the chiliensis group of species is thought to include almost half the species in genus.[7] It is hypothesized that chiliensis lizards originated from an Andean-Patagonean area.[4]

Description edit

 
An illustration of Calotes chilensis from 1700-1880[8]

The Chilean tree iguana is relatively large-sized compared to other lizards, reaching 25 cm (9.8 in) in total length. It is an oviparous species. It is carnivorous and feeds mainly on insects and other invertebrates.[9] The maximum body weight is 24.03g.[10] L. chiliensis individuals in more southern latitudes tend to be smaller in size than their northern latitude counterparts. Given their geographic distribution, lizards further south experience climates with lower temperatures.[11]

 
Photo of Lioaemus chilensis from Santiago, Chile in 2009
 
An illustration of Liolaemus chiliensis and Liolaemus cyanogaster
 
An illustration of Liolaemus chiliensis

These dull-colored lizards tend to be green and gray, helping them camouflage against rocks and vegetation. Axilla groin distance is higher in females than in males. Polyploidy results in morphological differences among lizards of different ploidy (i.e. in head shape and various proportions). The mechanisms behind these physical differences are unclear at this time.[12]

Morphology edit

Like most reptiles, this species has a tympanic membrane. Their otoliths help detect both acceleration and sound.[13] Their hearing range extends from 500 Hz to 2000 Hz, with a threshold of 40 dB.[14] These lizards excel at recognizing the distress calls of their own species.

Polyploidy edit

Polyploidy is rare in non-sterile reptiles, but is common in this species. L. chiliensis has populations with diploid (2n) individuals, triploid (3n) individuals, and diploid-triploid (2n/3n) mosaic lizards. One study found that[15] "33% of females were triploid, 57.1% were mosaics, and 9.5% were diploid" and "86% of L. chiliensis males were mosaics (2n/3n) and 14% were diploids."

Lizards may receive a reduced (n) or unreduced (2n) euploid gamete from their father.[16] In mosaic males, spermatogenesis derives from both diploid and triploid spermatogonia. Although polyploidy can result from interspecific hybridization, Raúl Araya-Donoso, Fernando Torres-Pérez, David Véliz, and Madeleine Lamborot found that a "relationship between hybridization and polyploidy could not be established because triploid and mosaic lizards were hybrids and purebreds."[17]

In general, polyploid individuals can be phenotypically different than diploid individuals. Studies have shown that polyploid individuals can have metabolic differences from diploid individuals, and may be more flexible in adapting to changing environments.[17]

Habitat and Distribution edit

This species is native to the Neotropics.[18] It is also found in the Chilean Matorral, Patagonian Steppe, and Valdivian Temperate Forests, covering a region spanning Chile and Argentina.[19] A helpful map of its distribution and numerous observations (and their photos) can be found on iNaturalist. This lizard lives within the latitudes of 30 and 40°S, and its altitudes are from sea level to 2100 meters.[12]

The habitat of this species includes bushes and branches. These lizards bask in bushes that tend to obscure them from possible predators, which makes it more difficult for predators to spot them. L. chiliensis is adept in running and climbing, and can often be found on branches higher than expected.[20] However, they are typically ground dwellers, found within dense shrubs but not on the ground directly.[21] Their geographical distribution extends from Coquimbo (29°S) to Valdivia (39°S).[22] L. chiliensis is active throughout the day. Observations tend to be more frequent from September to February, which may be due to its mating season, the warmer weather in these months, or human beings going out more often in warmer weather and consequentially spotting more of these lizards.

Behavior edit

Diet edit

The diet of L. chiliensis consists mostly of insects and small invertebrates. Juveniles behave and hunt similarly to adults. Studies in species within the chiliensis group have identified that these lizards feed mainly on Coleoptera, Hemiptera, and Hymenoptera. Stomach and fecal contents also included parts of grasshoppers, scorpions, plants, Lepidoptera, and Diptera.[23]

Eye Bulging edit

These lizards have been observed to display eye-bulging, which is thought to serve a purpose of thermoregulation or even eye-cleaning.[24] This behavior has been observed rarely and only in captivity.

Reproduction and Life Cycle edit

The weeping lizard is oviparous which means that female lizards lay eggs that hatch outside of her after a period of incubation without her presence. Females prefer to lay their eggs underneath rocks, where the eggs gain protection from predators. Females can lay their eggs as early as October, but typically, females are gravid in October and lay eggs in November shortly after the beginning of the warmer season in the Southern hemisphere. Clutches of eggs are usually between 7 and 11 eggs, laid together.[25] The maximum clutch size ever recorded has been 18 eggs.[26] Significant variation in egg clutch size has been documented before. Generally, larger females tend to lay more eggs than smaller females.[22] This trend applies to both mass and volume, since more massive lizards are almost always more voluminous. As a result, previous variation in egg-laying clutch size may actually be a result of unconsidered variation in the sizes of the females who laid the clutches.

Distress Calls edit

Vocal sound production is rare in lizards outside of Gekkota. The vast majority of non-Gekkota lizards are voiceless, with the exception of the weeping lizard, which is the only species in this 'voiceless' genus that can vocalize.[27] Weeping lizards are known to produce "distress calls" when stressed, cornered, or threatened, including when seized by humans. Both males and females produces these calls, which are known to sound more like squeaks than hisses or weeping. Thought to be produced by the larynx,[28] these sounds extend into the ultrasonic range. These distress calls can be heard several meters away from the lizard producing them.[28] These calls increase fear in other lizards. When scared, L. chiliensis become immobile and delay any attempts to escape.[29] The presence of a somewhat consistently high risk of predation reduces the activity of these lizards over long-term observation.[30]

Distress calls can vary in frequency, linearity, and the presence of harmonics. Compared to females, males tend to emit calls that are louder, more linear, and less complicated. The vocalizations are thought to serve two purposes: 1) they cause antipredatory behavior in conspecific neighbors and 2) possibly deter predators. Complex (nonlinear, erratic) calls have been shown to induce more fear in conspecific lizards than simple calls; the producer of the call may have less control over his or her call if there is a more alarming threat. Distress calls are also more distressed when heard in a familiar environment. Lizards may feel safer in a familiar environment, and therefore more alarmed by surprising causes of distress. On the other hand, the distress calls have been shown to reduce snake predation via a temporary reduction in their exploration, although there is a risk that the vocalization helps a predator narrow down the vocalizer's location.[31] Although seemingly counterintuitive, attracting multiple predators with a vocalization would also serve to be advantageous, because the larger predators would prioritize its safety and the possible conflict with approaching secondary or primary predators in the proximity.[32] When L. chiliensis could also detect the chemical scents of other weeping lizards, they mobilized more quickly and more often compared to without the scent. Without the scent, lizards froze to avoid detection; when other members of the same species were thought to be around, they were quicker to try escaping. Distress calls therefore have been evolutionarily advantageous, increasing the chances of survival both with or without other lizards present.

Members of L. chiliensis from different regions have also shown the ability to distinguish calls from individuals of their own region. Individual L. chiliensis in the northern area tend to react only to distress calls from other northern lizards, whereas southern lizards respond to distress calls from both northern and southern lizards.[11] Given that southern lizards tend to be smaller in size, their perceived risk of predation may be higher, causing increased sensitivity to distress calls produced by weeping lizards of any region. More research is needed into their means of evasion and escape.

While these distress calls can serve many advantageous purposes for the weeping lizard, there is evidence that the species Liolaemus lemniscatus will eavesdrop of these distress calls. This ability allows the Liolaemus lemniscatus to better escape its weeping lizard predator because the distress calls can serve as information of predation risk to the prey of weeping lizards. Therefore, these distress calls might negatively impact the weeping lizards’ ability to capture prey.[33]

Intraspecific Interactions edit

A study from the Journal of Herpetology suggests that L. chiliensis can detect other members of their species via the presence of lipids in recently deposited feces; however, there is no evidence of further recognition beyond the knowledge that another member of the same species is nearby.[34] Both male and females can detect female precloacal secretions of fellow L. chiliensis.[35] Once detected, these female secretions prompt exploration and movement of both sexes. Higher interest and exploration are exhibited in males that are possibly seeking a mate. Similarly, females may benefit from releasing these secretions by attracting potential mates.

Interspecific Interactions edit

L. chiliensis is sometimes preyed upon by the Chilean Green Racer (Philodryas chamissonis).[18] Spauligodon, a genus of host-specific nematodes, is a parasite towards these lizards. Liolaemus lizards are the most preyed upon lizard in their native Chilean habitat, due to their abundance and ecological niche.[21] Furthermore, these lizards are capable of detecting predatory snakes through scent. Detection resulted in a corresponding decrease in activity from the lizard.[36]

Conservation edit

L. chiliensis are kept as pets,[37] although such a practice violates conservation biology principles, because of the limited range of L. Chiliensis, and is thus vulnerable to habitat disruption and population decline. The species has no history of endangerment or near extinction; today, it is a species of least concern. Their popularity as a pet remains, due to their common presence within this range and their distress calls. These lizards have been photographed being held in human hands near its habitat in natural terrain, suggesting either that a human was able to quickly seize a cautious lizard or that the lizard was curious and calm enough to let itself be held.

References edit

  1. ^ THE REPTILE DATABASE. Liolaemus chiliensis (LESSON, 1830)
  2. ^ AuthorsAndGroup. Year. Talcahuano Smooth-throated Lizard Liolaemus chiliensis
  3. ^ C. Michael Hogan & World Wildlife Fund. 2013. Chilean matorral. ed. M.McGinley. Encyclopedia of Earth. National Council for Science and the Environment. Washington DC
  4. ^ a b c Juan Manuel Díaz GómezFernando Lobo. 2006. Historical Biogeography of a Clade of Liolaemus (Iguania: Liolaemidae) based on ancestral areas and dispersal-vicariance analysis (DIVA)
  5. ^ Lobo Gaviola, Fernando Jose. 2001. A phylogenetic analysis of lizards of the Liolaemus chiliensis group (lguania Tropiduridae)
  6. ^ Andrés Sebastián Quinteros, Cristian Simón Abdala, Juan Manuel Díaz Gómez, Gustavo José Scrocchi. 2008. Two New Species of Liolaemus (Iguania: Liolaemidae) of Central West Argentina
  7. ^ FERNANDO LOBO. 2001. A PHYLOGENETIC ANALYSIS OF LIZARDS OF THE LIOLAEMUS CHILIENSIS GROUP (IGUANIA: TROPIDURIDAE). HERPETOLOGICAL JOURNAL, Vol. 1 1 , pp. 1 37-1 50.
  8. ^ Iconographia Zoologica - Special Collections University of Amsterdam. 1700-1880. Calotes chilensis - 1700-1880
  9. ^ "Liolaemus chiliensis". Archived from the original on 2013-12-02. Retrieved 2013-11-26.
  10. ^ Hone et al. 2013. Body Size datasets for PLOS ONE paper
  11. ^ a b Antonieta Labra, Claudio Reyes-Olivares, Michael Weymann. Asymmetric Response to Heterotypic Distress Calls in the Lizard Liolaemus chiliensis.
  12. ^ a b Raúl Araya-Donoso, David Véliz, Marcela Vidal, and Madeleine Lamborot. 2017. Relationships of the morphological variation in diploids, triploids and mosaics of Liolaemus chiliensis (Sauria: Liolaemidae)
  13. ^ Ladich and Popper. 2004. Parallel Evolution in Fish Hearing Organs. Springer Handbook of Auditory Research, 95–127
  14. ^ Dooling et al. 2000. Hearing in Birds and Reptiles. Comparative Hearing: Birds and Reptiles
  15. ^ Madeleine Lamborot, M E Manzur, and E Alvarez-Sarret. 2006. Triploidy and mosaicism in Liolaemus chiliensis (Sauria: Tropiduridae)
  16. ^ Raúl Araya-Donoso, David Véliz, Marcela Vidal, and Madeleine Lamborot. 2017. Relationships of the morphological variation in diploids, triploids and mosaics of Liolaemus chiliensis (Sauria: Liolaemidae)
  17. ^ a b Raúl Araya-Donoso, Fernando Torres-Pérez, David Véliz, Madeleine Lamborot. 2019. Hybridization and polyploidy in the weeping lizard Liolaemus chiliensis (Squamata: Liolaemidae)
  18. ^ a b "Chilean Tree Iguana". EOL. Retrieved 7 October 2021.
  19. ^ World Wildlife Fund. 2006. [www.worldwildlife.org/WildFinder Online database of species distributions, ver. Jan-06]
  20. ^ Donoso-Barros, R. 1966. Reptiles de Chile. Santiago, Chile. Ediciones de la Universidad de Chile
  21. ^ a b Jaksić, Fabian M.; Greene, Harry W.; Schwenk, Kurt; Seib, Robert L. (January 1982). "Predation upon reptiles in Mediterranean habitats of Chile, Spain and California: A comparative analysis". Oecologia. 53 (2): 152–159. Bibcode:1982Oecol..53..152J. doi:10.1007/BF00545658. PMID 28311104. S2CID 22881961. Retrieved 28 November 2021.
  22. ^ a b Jaime Troncoso-Palacios, and Antonieta Labra. 2017. New egg laying record for Liolaemus chiliensis (Lesson, 1830) (Iguania: Liolaemidae)
  23. ^ Halloy1, Robles2, Cuezzo3, Monique1, Cecilia2, Fabiana3 (2006). "Diet in two syntopic neotropical lizard species of Liolaemus (Liolaemidae): interspecific and intersexual differences". Rev. Esp. Herp. 20: 47–56.{{cite journal}}: CS1 maint: multiple names: authors list (link) CS1 maint: numeric names: authors list (link)
  24. ^ Reyes Olivares, Claudio; Rain Garrido, Inger; Labra, Antonieta. 2016. The eye-bulging in Liolaemus lizards (Weigmann 1843)
  25. ^ Pincheira-Donoso, D., Núñez, H. 2005. Les especies Chilenas del género Liolaemus Wiegmann, 1834 (Iguania: Tropiduridae: Liolaeminae): taxonomía, sistemática y evolución. Museo Nacional de Historia Natural Chile
  26. ^ Ibargüengoytía. 2008. Estrategias reproductivas en reptiles.
  27. ^ Antonieta Labra, Gabriela Silva, Fernanda Norambuena, Nelson Velásquez and Mario Penna. 2013. Acoustic Features of the Weeping Lizard's Distress Call
  28. ^ a b John H. Carothers, Jeffrey G. Groth & Fabian M. Jaksic. 2010. Vocalization as a Response to Capture in the Central Chilean Lizard Liolaemus chiliensis (Tropiduridae)
  29. ^ Mario R. Ruiz-Monachesi and Antonieta Labra. Complex distress calls sound frightening: the case of the weeping lizard.
  30. ^ Javiera Constanzo-Chávez, Mario Penna, Antonieta Labra. 2018. Comparing the antipredator behaviour of two sympatric, but not syntopic, Liolaemus lizards
  31. ^ Misque Hoare, Antonieta Labra. 2013. Searching for the Audience of the Weeping Lizard's Distress Call
  32. ^ Högstedt. 1983. Snake modulates constriction in response to prey's heartbeat
  33. ^ Fong, Lydia J. M.; Navea, Fernando; Labra, Antonieta (2021). "Does Liolaemus lemniscatus eavesdrop on the distress calls of the sympatric weeping lizard?". Journal of Ethology. 39. Lydia J. M. Fong: 11–17. doi:10.1007/s10164-020-00666-y. S2CID 220966872.
  34. ^ Soledad Valdecantos; Mario R. Ruiz-Monachesi; Antonieta Labra. 2020. Testing the Functionality of Lipids from Feces in the Conspecific Recognition of the Weeping Lizard, Liolaemus chiliensis
  35. ^ Soledad Valdecantos, Antonieta Labra. 2017. Testing the functionality of precloacal secretions from both sexes in the South American lizard, Liolaemus chiliensis
  36. ^ Labra, Antonieta; Hoare, Misque (17 September 2014). "Chemical recognition in a snake–lizard predator–prey system". Acta Ethologica. 18 (2): 173–179. doi:10.1007/s10211-014-0203-7. S2CID 14953912. Retrieved 28 November 2021.
  37. ^ Herman A.J. in den Bosch. "How to keep lacertids: Liolaemus". Podarcis. Retrieved 2012-09-19.
  • Podarcis: How to keep lizards
  • Mi Fauna: Liolaemus chiliensis