Animals maintain complex associations with a diverse microbiota living in their guts. Our understanding of the ecology of these associations is extremely limited in reptiles. Here, we report an in-depth study into the microbial ecology of gut communities in three syntopic and viviparous lizard species (two omnivores: Liolaemus parvus and Liolaemus ruibali and an herbivore: Phymaturus williamsi). Using 16S rRNA gene sequencing to inventory various bacterial communities, we elucidate four major findings: (i) closely related lizard species harbour distinct gut bacterial microbiota that remain distinguishable in captivity; a considerable portion of gut bacterial diversity (39.1%) in nature overlap with that found on plant material, (ii) captivity changes bacterial community composition, although host-specific communities are retained, (iii) faecal samples are largely representative of the hindgut bacterial community and thus represent acceptable sources for nondestructive sampling, and (iv) lizards born in captivity and separated from their mothers within 24 h shared 34.3% of their gut bacterial diversity with their mothers, suggestive of maternal or environmental transmission. Each of these findings represents the first time such a topic has been investigated in lizard hosts. Taken together, our findings provide a foundation for comparative analyses of the faecal and gastrointestinal microbiota of reptile hosts.
While herbivory is a common feeding strategy in a number of vertebrate classes, less than 4% of squamate reptiles feed primarily on plant material. It has been hypothesized that physiological or microbial limitations may constrain the evolution of herbivory in lizards. Herbivorous lizards exhibit adaptations in digestive morphology and function that allow them to better assimilate plant material. However, it is unknown whether these traits are fixed or perhaps phenotypically flexible as a result of diet. Here, we maintained a naturally omnivorous lizard, Liolaemus ruibali, on a mixed diet of 50% insects and 50% plant material, or a plant-rich diet of 90% plant material. We compared parameters of digestive performance, gut morphology and function, and gut microbial community structure between the two groups. We found that lizards fed the plant-rich diet maintained nitrogen balance and exhibited low minimum nitrogen requirements. Additionally, lizards fed the plantrich diet exhibited significantly longer small intestines and larger hindguts, demonstrating that gut morphology is phenotypically flexible. Lizards fed the plant-rich diet harbored small intestinal communities that were more diverse and enriched in Melainabacteria and Oscillospira compared with mixed diet-fed lizards. Additionally, the relative abundance of sulfate-reducing bacteria in the small intestine significantly correlated with whole-animal fiber digestibility. Thus, we suggest that physiological and microbial limitations do not sensu stricto constrain the evolution of herbivory in lizards. Rather, ecological context and fitness consequences may be more important in driving the evolution of this feeding strategy.
The diversity of habitats generated by the Andes uplift resulted a mosaic of heterogeneous environments in South America for species to evolve a variety of ecological and physiological specializations. Species in the lizard family Liolaemidae occupy a myriad of habitats in the Andes. Here, we analyze the tempo and mode of evolution in the thermal biology of liolaemids. We assessed whether there is evidence of local adaptation (lability) or conservatism (stasis) in thermal traits. We tested the hypothesis that abiotic factors (e.g., geography, climate) rather than intrinsic factors (egg‐laying [oviparous] or live‐bearing [viviparous], substrate affinity) explain variation in field active body temperature (Tb), preferred temperature (Tp), hours of restriction of activity, and potential hours of activity. Although most traits exhibited high phylogenetic signal, we found variation in thermal biology was shaped by geography, climate, and ecological diversity. Ancestral character reconstruction showed shifts in Tb tracked environmental change in the past ∼20,000 years. Thermal preference is 3°C higher than Tb, yet exhibited a lower rate of evolution than Tb and air temperature. Viviparous Liolaemus have lower Tbs than oviparous species, whereas Tp is high for both modes of reproduction, a key difference that results in a thermal buffer for viviparous species to cope with global warming. The rapid increase in environmental temperatures expected in the next 50–80 years in combination with anthropogenic loss of habitats are projected to cause extirpations and extinctions in oviparous species.
The South American lizard genus Liolaemus comprises > 260 species, of which > 60 are recognized as members of the Liolaemus montanus group, distributed throughout the Andes in central Peru, Bolivia, Chile and central Argentina. Despite its great morphological diversity and complex taxonomic history, a robust phylogenetic estimate is still lacking for this group. Here, we study the morphological and molecular diversity of the L. montanus group and present the most complete quantitative phylogenetic hypothesis for the group to date. Our phylogeny includes 103 terminal taxa, of which 91 are members of the L. montanus group (58 are assigned to available species and 33 are of uncertain taxonomic status). Our matrix includes 306 morphological and ecological characters and 3057 molecular characters. Morphological characters include 48 continuous and 258 discrete characters, of which 70% (216) are new to the literature. The molecular characters represent five mitochondrial markers. We performed three analyses: a morphology-only matrix, a molecular-only matrix and a matrix including both morphological and molecular characters (total evidence hypothesis). Our total evidence hypothesis recovered the L. montanus group as monophyletic and included ≥ 12 major clades, revealing an unexpectedly complex phylogeny.
In ectotherms, temperature exerts a strong influence on the performance of physiological and ecological traits. One approach to understand the impact of rising temperatures on animals and their ability to cope with climate change is to quantify variation in thermal-sensitive traits. Here, we examined the thermal biology, the temperature dependence and the thermal plasticity of bite force (endurance and magnitude) in Diplolaemus leopardinus, an aggressive and territorial lizard, endemic to Mendoza province, Argentina. Our results indicated that this lizard behaves like a moderate thermoregulator which uses the rocks of its environment as the main heat source. Bite endurance was not influenced by head morphometry and body temperature, whereas bite force was influenced by head length and jaw length, and exhibited thermal dependence. Before thermal acclimation treatments, the maximum bite force for D. leopardinus occured at the lowest body temperature and fell sharply with increasing body temperature. After acclimation treatments, lizards acclimated at higher temperatures exhibited greater bite force. Bite force showed phenotypic plasticity, which reveals that leopard iguanas are able to maintain (and even improve) their bite force under a rising-temperature scenario.
As a result of several field trips and studies of collections of Phymaturus samples from Andean areas of central western Argentina (San Juan province), we discovered two populations that exhibit a particular character combination not seen in other species formally recognized in the literature. Based on a detailed analysis of an extended list of morphological characters (93), including scalation, color pattern, gular and nuchal folds, precloacal pores, and morphometric data, we conclude that these populations represent independent lineages that deserve to be considered as new species. According to the most recent revision of the genus and considering the descriptions made in another recent contribution, the taxo-nomic composition of the genus was increased to 38 species. In this study we provide the formal description of two addi-tional new taxa, including their diagnosis and detailed comparisons with other members of their species group. The two new species belong to the palluma group, and can be assigned to the Puna subclade because they present the typical dorsal “spray” pattern. Other characters described in this study suggest their close phylogenetic relationship with other species of this subclade inhabiting the southern Puna region of Argentina, such as Phymaturus punae. Within the Puna subclade, Phymaturus aguanegra sp. nov. differs from all other members (P. antofagastensis, P. denotatus, P. laurenti, P. punae, P. extrilidus, P. mallimaccii and P. paihuanense) exhibiting the following combination of diagnostic characters: a complete melanism over the dorsum of neck, the presence of enlarged scales at the base of tail in males, having strongly keeled tarsal scales, lacking enlarged scales on the anterior margin of the antehumeral fold and centre of chest, females without flank coloration, a vertebral dark gray stripe usually present on the dorsum, females exhibiting a tricolor dorsal pattern, with two types of brown and scattered ferriferous oxide spots, and the absence of a scapular spot. Phymaturus williamsi sp. nov. differs from all other members of the Puna clade because: exhibits an “aggregate” dorsal pattern, unlike the ho-mogeneous spray of most Puna species, lacks enlarged scales on the anterior margin of the antehumeral fold and in the centre of chest, flank coloration in females is absent, females of Phymaturus williamsi sp. nov. lack white transversal stripes on the dorsal pattern, preocular scale in contact with canthal scale in Phymaturus williamsi sp. nov., rostral scale can be divided in Phymaturus williamsi sp. nov. and shows the largest number of scales counted around midbody within the Puna subclade (x= 213.4; 186-235).
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