Liolaemus chiliensis

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.{{cite journal | last1=Díaz Gómez | first1=Juan Manuel | last2=Lobo | first2=Fernando | title=Historical Biogeography of a Clade of Liolaemus (Iguania: Liolaemidae) based on ancestral areas and dispersal-vicariance analysis (DIVA) | journal=Papéis Avulsos de Zoologia (São Paulo) | publisher=Universidade de Sao Paulo, Agencia USP de Gestao da Informacao Academica (AGUIA) | volume=46 | issue=24 | year=2006 | issn=0031-1049 | doi=10.1590/s0031-10492006002400001 | doi-access=free | url=https://www.scielo.br/j/paz/a/q5KtqfwhN3wRGLTjbYq4SnB/?lang=en&format=pdf }} 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. 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.{{cite journal | last1=Gaviola | first1=Lobo | last2=Jose | first2=Fernando | title=A phylogenetic analysis of lizards of the Liolaemus chiliensis group (lguania Tropiduridae) |journal=Herpetological Journal | publisher=British Herpetological Society | date=December 2001 |pages=137–150 | issn=0268-0130 | url=https://ri.conicet.gov.ar/handle/11336/54988 }} Some have distinguished multiple species within L. chiliensis due to differences in "snout-vent length, squamation, and dorsal and ventral colors."{{cite journal | last1=Quinteros | first1=Andrés Sebastián | last2=Abdala | first2=Cristian Simón | last3=Díaz Gómez | first3=Juan Manuel | last4=Scrocchi | first4=Gustavo José | title=Two New Species of Liolaemus (Iguania: Liolaemidae) of Central West Argentina | journal=South American Journal of Herpetology | volume=3 | issue=2 | date=2008 | issn=1808-9798 | doi=10.2994/1808-9798(2008)3[101:TNSOLI]2.0.CO;2 | doi-access=free | pages=101–111 | hdl=11336/97456 | url=https://ri.conicet.gov.ar/bitstream/11336/97456/2/CONICET_Digital_Nro.9159b440-3fd9-4c53-ba90-b610f36a651e_A.pdf }} 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. It is hypothesized that chiliensis lizards originated from an Andean-Patagonean area.

File:Calotes chilensis - 1700-1880 - Print - Iconographia Zoologica - Special Collections University of Amsterdam - UBA01 IZ12800117.tif

The Chilean tree iguana is relatively large-sized compared to other lizards, reaching {{convert|25|cm}} in total length. It is an oviparous species. It is carnivorous and feeds mainly on insects and other invertebrates.{{Cite web |url=http://www.atlasherpetozoos.cl/Reptiles/LChiliensis.html |title=Liolaemus chiliensis |access-date=2013-11-26 |archive-url=https://web.archive.org/web/20131202221557/http://www.atlasherpetozoos.cl/Reptiles/LChiliensis.html |archive-date=2013-12-02 |url-status=dead }} The maximum body weight is 24.03g.{{citation | last1=Hone | first1=David | last2=O'Gorman | first2=Eoin J. | title=Body Size datasets for PLOS ONE paper doi:10.1371 | date=2013 | publisher=Figshare | doi=10.6084/M9.FIGSHARE.627530 | doi-access=free | url=https://figshare.com/articles/dataset/Body_Size_datasets_for_PLOS_ONE_paper_doi_10_1371/627530 }} 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.{{cite journal | last1=Labra | first1=Antonieta | last2=Reyes-Olivares | first2=Claudio | last3=Weymann | first3=Michael | title=Asymmetric Response to Heterotypic Distress Calls in the Lizard Liolaemus chiliensis | journal=Ethology | volume=122 | issue=9 | date=2016 | issn=0179-1613 | doi=10.1111/eth.12522 | pages=758–768 | bibcode=2016Ethol.122..758L | url=https://onlinelibrary.wiley.com/doi/10.1111/eth.12522 }}

File:Lizard-santiago-2009-17-01.JPG

File:Liolaemus chiliensis Liolaemus cyanogaster.JPG

File:Liolaemus chiliensis.jpg

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.{{cite journal | last1=Araya-Donoso | first1=Raúl | last2=Véliz | first2=David | last3=Vidal | first3=Marcela | last4=Lamborot | first4=Madeleine | title=Relationships of the morphological variation in diploids, triploids and mosaics of Liolaemus chiliensis (Sauria: Liolaemidae) | journal=Amphibia-Reptilia | publisher=Walter de Gruyter GmbH | volume=38 | issue=4 | year=2017 | issn=0173-5373 | doi=10.1163/15685381-00003132 | pages=503–515}}

Like most reptiles, this species has a tympanic membrane. Their otoliths help detect both acceleration and sound.{{cite book | last1=Ladich | first1=Friedrich | last2=Popper | first2=Arthur N. | title=Springer Handbook of Auditory Research | chapter=Parallel Evolution in Fish Hearing Organs | publisher=Springer New York | publication-place=New York, NY | year=2004 | isbn=978-0-387-21093-3 | issn=0947-2657 | doi=10.1007/978-1-4419-8957-4_4 | pages=95–127}} Their hearing range extends from 500 Hz to 2000 Hz, with a threshold of 40 dB.{{cite book | last1=Dooling | first1=Robert J. | last2=Lohr | first2=Bernard | last3=Dent | first3=Micheal L. | title=Comparative Hearing: Birds and Reptiles | chapter=Hearing in Birds and Reptiles | publisher=Springer New York | publication-place=New York, NY | volume=13 | date=2000 | isbn=978-1-4612-7036-2 | doi=10.1007/978-1-4612-1182-2_7 | chapter-url=http://link.springer.com/10.1007/978-1-4612-1182-2_7 | pages=308–359}} These lizards excel at recognizing the distress calls of their own species.

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{{cite journal | last1=Lamborot | first1=Madeleine | last2=Manzur | first2=M E | last3=Alvarez-Sarret | first3=E | title=Triploidy and mosaicism in Liolaemus chiliensis (Sauria: Tropiduridae) | journal=Genome | volume=49 | issue=5 | date=2006-05-01 | issn=0831-2796 | doi=10.1139/g05-122 | pages=445–453 | pmid=16767169 | url=http://www.nrcresearchpress.com/doi/10.1139/g05-122 }} "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. 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."{{cite journal | last1=Araya-Donoso | first1=Raúl | last2=Torres-Pérez | first2=Fernando | last3=Véliz | first3=David | last4=Lamborot | first4=Madeleine | title=Hybridization and polyploidy in the weeping lizard Liolaemus chiliensis (Squamata: Liolaemidae) | journal=Biological Journal of the Linnean Society | date=2019-10-23 | issn=0024-4066 | doi=10.1093/biolinnean/blz145 | url=https://academic.oup.com/biolinnean/advance-article/doi/10.1093/biolinnean/blz145/5603699 }}

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.

This species is native to the Neotropics. It is also found in the Chilean Matorral, Patagonian Steppe, and Valdivian Temperate Forests, covering a region spanning Chile and Argentina. This lizard lives within the latitudes of 30 and 40°S, and its altitudes are from sea level to 2100 meters.

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.{{cite book |last=Donoso-Barros |first=Roberto |year=1966 |url=https://openlibrary.org/books/OL5651016M/Reptiles_de_Chile |title=Reptiles de Chile |location=Santiago, Chile |publisher=Ediciones de la Universidad de Chile|ol=5651016M }} However, they are typically ground dwellers, found within dense shrubs but not on the ground directly.{{cite journal|last1=Jaksić|first1=Fabian M.|last2=Greene|first2=Harry W.|last3=Schwenk|first3=Kurt|last4=Seib|first4=Robert L.|date=January 1982|title=Predation upon reptiles in Mediterranean habitats of Chile, Spain and California: A comparative analysis|url=https://link.springer.com/article/10.1007/BF00545658|journal=Oecologia|volume=53|issue=2|pages=152–159|doi=10.1007/BF00545658|pmid=28311104|bibcode=1982Oecol..53..152J|s2cid=22881961|access-date=28 November 2021}} Their geographical distribution extends from Coquimbo (29°S) to Valdivia (39°S)．{{cite journal | last1=Troncoso-Palacios | first1=Jaime | last2=Labra | first2=Antonieta | title=New egg laying record for Liolaemus chiliensis (Lesson, 1830) (Iguania: Liolaemidae) | journal=Herpetology Notes | volume=10 | date=2017-10-10 | issn=2071-5773 | pages=529–531 | url=https://www.biotaxa.org/hn/article/view/30658 }} 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.

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.{{Cite journal| last1=Halloy | first1=Monique | last2=Robles | first2=Cecilia Ines | last3=Cuezzo | first3=Fabiana del Carmen|date=2006|title=Diet in two syntopic neotropical lizard species of Liolaemus (Liolaemidae): interspecific and intersexual differences|url=https://www.academia.edu/18199181|journal=Rev. Esp. Herp.|volume=20|pages=47–56}}

These lizards have been observed to display eye-bulging, which is thought to serve a purpose of thermoregulation or even eye-cleaning.{{cite journal | last1=Olivares | first1=Reyes | last2=Garrido | first2=Rain | last3=Labra | first3=Antonieta | title=The eye-bulging in Liolaemus lizards (Weigmann 1843) |journal=Gayana |volume=80 |issue=1 |pages=129–132 | date=2016 | doi=10.4067/S0717-65382016000100013 | issn=0717-6538 | url=https://repositorio.uchile.cl/handle/2250/142881 }} This behavior has been observed rarely and only in captivity.

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.{{cite journal | last1=Pincheira-Donoso | first1=Daniel | last2=Nuñez | first2=Herman | title=Las especies chilenas del genero Liolaemus Wiegmann, 1834 (Iguania: Tropiduridae: Liolaeminae): Taxonomía, sistemática y evolución | journal=Publicaciones Ocasionales Museo Nacional de Historia Natural de Chile | volume=59 | date=2005 | issn=0716-0224 | pages=1–486 | url=https://pure.qub.ac.uk/en/publications/las-especies-chilenas-del-genero-liolaemus-wiegmann-1834-iguania- }} The maximum clutch size ever recorded has been 18 eggs.{{cite book|last=Ibargüengoytía |first= N. R. |year=2008 |chapter=Estrategias reproductivas en reptiles |title=Herpetología de Chile |pages=391–425}} Significant variation in egg clutch size has been documented before. Generally, larger females tend to lay more eggs than smaller females. 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.

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.{{cite journal | last1=Labra | first1=Antonieta | last2=Silva | first2=Gabriela | last3=Norambuena | first3=Fernanda | last4=Velásquez | first4=Nelson | last5=Penna | first5=Mario | title=Acoustic Features of the Weeping Lizard's Distress Call | journal=Copeia | publisher=[American Society of Ichthyologists and Herpetologists (ASIH), Allen Press] | volume=2013 | issue=2 | year=2013 | issn=0045-8511 | jstor=24635712 | pages=206–212 | doi=10.1643/CE-12-026 | url=http://www.jstor.org/stable/24635712 }} 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,{{cite journal | last1=Carothers | first1=John H. | last2=Groth | first2=Jeffrey G. | last3=Jaksic | first3=Fabian M. | title=Vocalization as a Response to Capture in the Central Chilean Lizard Liolaemus chiliensis (Tropiduridae) | journal=Studies on Neotropical Fauna and Environment | volume=36 | issue=2 | date=2001 | issn=0165-0521 | doi=10.1076/snfe.36.2.93.2136 | pages=93–94 | bibcode=2001SNFE...36...93C | url=http://www.tandfonline.com/doi/abs/10.1076/snfe.36.2.93.2136 }} these sounds extend into the ultrasonic range. These distress calls can be heard several meters away from the lizard producing them. These calls increase fear in other lizards. When scared, L. chiliensis become immobile and delay any attempts to escape.{{cite journal | last1=Ruiz-Monachesi | first1=Mario R. | last2=Labra | first2=Antonieta | title=Complex distress calls sound frightening: the case of the weeping lizard | journal=Animal Behaviour | publisher=Elsevier BV | volume=165 | year=2020 | issn=0003-3472 | doi=10.1016/j.anbehav.2020.05.004 | doi-access=free | pages=71–77}} The presence of a somewhat consistently high risk of predation reduces the activity of these lizards over long-term observation.{{cite journal | last1=Constanzo-Chávez | first1=Javiera | last2=Penna | first2=Mario | last3=Labra | first3=Antonieta | title=Comparing the antipredator behaviour of two sympatric, but not syntopic, Liolaemus lizards | journal=Behavioural Processes | publisher=Elsevier BV | volume=148 | year=2018 | issn=0376-6357 | doi=10.1016/j.beproc.2018.01.005 | pages=34–40| pmid=29330087 }}

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.{{cite journal | last1=Hoare | first1=Misque | last2=Labra | first2=Antonieta | title=Searching for the Audience of the Weeping Lizard's Distress Call | journal=Ethology | volume=119 | issue=10 | date=2013 | issn=0179-1613 | doi=10.1111/eth.12127 | pages=860–868 | bibcode=2013Ethol.119..860H | url=https://onlinelibrary.wiley.com/doi/10.1111/eth.12127 }} 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.{{cite journal | last1=Boback | first1=Scott M. | last2=Hall | first2=Allison E. | last3=McCann | first3=Katelyn J. | last4=Hayes | first4=Amanda W. | last5=Forrester | first5=Jeffrey S. | last6=Zwemer | first6=Charles F. | title=Snake modulates constriction in response to prey's heartbeat | journal=Biology Letters | volume=8 | issue=3 | date=2012-06-23 | issn=1744-9561 | pmid=22258447 | pmc=3367750 | doi=10.1098/rsbl.2011.1105 | doi-access=free | pages=473–476 }} 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. 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.{{cite journal |title=Does Liolaemus lemniscatus eavesdrop on the distress calls of the sympatric weeping lizard? |journal=Journal of Ethology |year=2021 |publisher=Lydia J. M. Fong|doi=10.1007/s10164-020-00666-y |last1=Fong |first1=Lydia J. M. |last2=Navea |first2=Fernando |last3=Labra |first3=Antonieta |volume=39 |pages=11–17 |s2cid=220966872 |doi-access=free |hdl=10852/83945 |hdl-access=free }}

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.{{cite journal | last1=Valdecantos | first1=Soledad | last2=Ruiz-Monachesi | first2=Mario R. | last3=Labra | first3=Antonieta | title=Testing the Functionality of Lipids from Feces in the Conspecific Recognition of the Weeping Lizard, Liolaemus chiliensis | journal=Journal of Herpetology | publisher=Society for the Study of Amphibians and Reptiles | volume=54 | issue=4 | date=2020-12-16 | issn=0022-1511 | doi=10.1670/19-074 }} Both male and females can detect female precloacal secretions of fellow L. chiliensis.{{cite journal | last1=Valdecantos | first1=Soledad | last2=Labra | first2=Antonieta | title=Testing the functionality of precloacal secretions from both sexes in the South American lizard, Liolaemus chiliensis | journal=Amphibia-Reptilia | publisher=Walter de Gruyter GmbH | volume=38 | issue=2 | year=2017 | issn=0173-5373 | doi=10.1163/15685381-00003106 | doi-access=free | pages=209–216 | hdl=11336/49960 | url=https://ri.conicet.gov.ar/bitstream/11336/49960/2/CONICET_Digital_Nro.fb56b458-dde7-46be-88d1-e32c49e0c3f3_A.pdf }} 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.

L. chiliensis is sometimes preyed upon by the Chilean Green Racer (Philodryas chamissonis).{{cite web |title=Chilean Tree Iguana |url=https://eol.org/pages/791942 |website=EOL |access-date=7 October 2021}} 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. Furthermore, these lizards are capable of detecting predatory snakes through scent. Detection resulted in a corresponding decrease in activity from the lizard.{{cite journal |last1=Labra |first1=Antonieta |last2=Hoare |first2=Misque |title=Chemical recognition in a snake–lizard predator–prey system |journal=Acta Ethologica |date=17 September 2014 |volume=18 |issue=2 |pages=173–179 |doi=10.1007/s10211-014-0203-7 |s2cid=14953912 |url=https://link.springer.com/article/10.1007/s10211-014-0203-7 |access-date=28 November 2021}}

L. chiliensis are kept as pets,{{cite web |url=http://www.podarcis.nl/info/halsbandhaguk.php3 |title=How to keep lacertids: Liolaemus|accessdate=2012-09-19 |format= |work= Podarcis|author= Herman A.J. in den Bosch}} 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.

{{reflist}}

• [http://www.podarcis.nl/info/halsbandhaguk.php3 Podarcis: How to keep lizards]
• [http://mifauna.blogspot.com/2007/04/traro-o-carancho-cordillerano.html Mi Fauna: Liolaemus chiliensis]

{{Commons category|Liolaemus chiliensis|position=left}}

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chilensis

Category:Reptiles described in 1830

Category:Taxa named by René Lesson

Category:Lizards of South America

Category:Reptiles of Chile

Category:Reptiles of Argentina

Category:Fauna of the Chilean Matorral