Desert Adaptation Research Articles

Effect of water stress on locomotor performance in Pleurodema nebulosum (Anura: Leptodactylidae) a native frog from the Argentina Monte desert

Locomotion is a fundamental aspect in the life cycle of any organism, and plays a crucial role in survival, feeding, escaping from predators, territoriality, and reproduction. Locomotion in amphibian anurans includes swimming, walking, jumping, climbing, and gliding. Amphibians living in arid regions face a constant compromise between hydration balance and locomotor performance. We studied the effects of water stress on the locomotion of the frog Pleurodema nebulosum, a specialist from the Monte Desert. Locomotor performance was measured through different physical variables during the jump. The locomotor performance of P. nebulosum showed a significant decrease in the magnitudes associated with a decrease in body hydration. Our study demonstrated how a desert-adapted frog (P. nebulosum) showed changes in locomotion related to the loss of body water. Additionally, we observed an abrupt drop in locomotion with 15% desiccation, followed by a plateau where the evaluated parameters remained stable. Pleuroderma nebulosum showed a clear adaptation to extreme desiccation. The frogs are rarely found in optimal hydration conditions as the environment in which they develop is extremely stressful from a water perspective. The data evaluated here are the beginning of a deeper exploration of the desert adaptations of South American desert frogs.

Adaptions of hunter-gatherers before Younger Dryas in Gobi Desert, northwest China: Flexible technological strategies in Pigeon Mountain Locality 10

The role of Oasis-based economies was pivotal for Hunter-gatherers in the Gobi Desert following the Last Glacial Maximum. Despite this significance, there has been a lack of systematic technological studies on lithic assemblages to investigate the technological strategies employed by hunter-gatherers for desert adaptation. In this study, we conducted a lithic analysis based on new excavations at Pigeon Mountain Locality 10 (QG10), focusing on archaeological remains from CL3 and CL2 dating back to 13.7–11.7 ka cal BP. Our findings reveal that the lithic technology employed reflects a flexible strategy, characterized by expedient flake production, the versatility of flake cores, and a diverse array of microblade cores and tools. Notably, we observed an expedient organizational strategy in microblade production, involving minimal investment in pre-shaping microblade cores and reduced depletion in core rejuvenation. Additionally, there was a heightened use of local quartzite raw materials, indicating enhanced flexibility, possibly influenced by the relatively dry and cold environmental changes during that period.

Geometric morphometrics of silky pocket mice (Perognathus: Perognathinae: Rodentia) crania reveals new insights into their variation, evolution, and taxonomy

Abstract We used cranial geometric morphometric methods (GMM) to explore interspecific variation in Perognathus (silky pocket mice). We digitized 67 cranial landmarks on photographs of 305 adult voucher specimens (10 species and 33 subspecies, 121 localities). After summarizing variation patterns, we explored their association with biological, ecological, and climatic factors, and how (and why) shape evolved through time. Phenotypic clusters did not accord with phylogenetic clades, and we found instances of convergence, divergence, and increased rates of shape change. These are discussed in terms of shared selective pressures (e.g. habitat substrate and burrowing plus historic shifts in habitat/climate and desert adaptation). We also used variation patterns and phenetics to address ongoing taxonomic issues. While pocket mice are difficult to distinguish externally, GMM allowed us to precisely characterize interspecific cranial variation. Species significantly differed and could be classified based on their shape. We find P. alticola to be the most cranially distinct species and suggest raising the P. alticola + P. parvus (including P. mollipilosus) cluster/clade to subgenus status (=Cricetodipus). Cranial GMM found differences among taxa that seemed similar (i.e. cryptic) when previously explored using linear measurements (e.g. P. parvus vs. P. mollipilosus).

Evolution of a fatty acyl-CoA elongase underlies desert adaptation in Drosophila.

Traits that allow species to survive in extreme environments such as hot-arid deserts have independently evolved in multiple taxa. However, the genetic and evolutionary mechanisms underlying these traits have thus far not been elucidated. Here, we show that Drosophila mojavensis, a desert-adapted fruit fly species, has evolved high desiccation resistance by producing long-chain methyl-branched cuticular hydrocarbons (mbCHCs) that contribute to a cuticular lipid layer reducing water loss. We show that the ability to synthesize these longer mbCHCs is due to evolutionary changes in a fatty acyl-CoA elongase (mElo). mElo knockout in D. mojavensis led to loss of longer mbCHCs and reduction of desiccation resistance at high temperatures but did not affect mortality at either high temperatures or desiccating conditions individually. Phylogenetic analysis showed that mElo is a Drosophila-specific gene, suggesting that while the physiological mechanisms underlying desert adaptation may be similar between species, the genes involved in these mechanisms may be species or lineage specific.

Whole-genome sequencing reveals adaptations of hairy-footed jerboas (Dipus, Dipodidae) to diverse desert environments

BackgroundEnvironmental conditions vary among deserts across the world, spanning from hyper-arid to high-elevation deserts. However, prior genomic studies on desert adaptation have focused on desert and non-desert comparisons overlooking the complexity of conditions within deserts. Focusing on the adaptation mechanisms to diverse desert environments will advance our understanding of how species adapt to extreme desert environments. The hairy-footed jerboas are well adapted to diverse desert environments, inhabiting high-altitude arid regions, hyper-arid deserts, and semi-deserts, but the genetic basis of their adaptation to different deserts remains unknown.ResultsHere, we sequenced the whole genome of 83 hairy-footed jerboas from distinct desert zones in China to assess how they responded under contrasting conditions. Population genomics analyses reveal the existence of three species in hairy-footed jerboas distributed in China: Dipus deasyi, Dipus sagitta, and Dipus sowerbyi. Analyses of selection between high-altitude desert (elevation ≥ 3000m) and low-altitude desert (< 500m) populations identified two strongly selected genes, ATR and HIF1AN, associated with intense UV radiation and hypoxia in high-altitude environments. A number of candidate genes involved in energy and water homeostasis were detected in the comparative genomic analyses of hyper-arid desert (average annual precipitation < 70mm) and arid desert (< 200mm) populations versus semi-desert (> 360mm) populations. Hyper-arid desert animals also exhibited stronger adaptive selection in energy homeostasis, suggesting water and resource scarcity may be the main drivers of desert adaptation in hairy-footed jerboas.ConclusionsOur study challenges the view of deserts as homogeneous environments and shows that distinct genomic adaptations can be found among desert animals depending on their habitats.

Review On the Desert Adaptations In Camel At Region Of Thar Desert In Churu District, Rajasthan, India

The study conducted in the Thar Region of Churu district including villages also at Rajasthan state in India. It includes a study on Desert adaptations which founds in Camel in Thar Desert. Thar desert one of the tropical deserts. In region of Thar Desert xerophytic plants and xerocole animals are found. All herbivorous animals depend on vegetations found here. Camel belongs to mammalia class of chordate phylum. The Camel is able to live in Thar Desert with its adaptations for this type of environment. Adaptations makes comfortable to an animal or a plant to well develop in particular area where they are founds. Find out the amazing adaptive features of camel in this study.

Similar adaptative mechanism but divergent demographic history of four sympatric desert rodents in Eurasian inland

Phenotypes associated with metabolism and water retention are thought to be key to the adaptation of desert species. However, knowledge on the genetic changes and selective regimes on the similar and divergent ways to desert adaptation in sympatric and phylogenetically close desert organisms remains limited. Here, we generate a chromosome level genome assembly for Northern three-toed jerboa (Dipus sagitta) and three other high-quality genome assemblies for Siberian jerboa (Orientallactaga sibirica), Midday jird (Meriones meridianus), and Desert hamster (Phodopus roborovskii). Genomic analyses unveil that desert adaptation of the four species mainly result from similar metabolic pathways, such as arachidonic acid metabolism, thermogenesis, oxidative phosphorylation, insulin related pathway, DNA repair and protein synthesis and degradation. However, the specific evolved genes in the same adaptative molecular pathway often differ in the four species. We also reveal similar niche selection but different demographic histories and sensitivity to climate changes, which may be related to the diversified genomic adaptative features. In addition, our study suggests that nocturnal rodents have evolved some specific adaptative mechanism to desert environments compared to large desert animals. Our genomic resources will provide an important foundation for further research on desert genetic adaptations.

Shared Patterns of Gene Expression and Protein Evolution Associated with Adaptation to Desert Environments in Rodents.

Desert specialization has arisen multiple times across rodents and is often associated with a suite of convergent phenotypes, including modification of the kidneys to mitigate water loss. However, the extent to which phenotypic convergence in desert rodents is mirrored at the molecular level is unknown. Here, we sequenced kidney mRNA and assembled transcriptomes for three pairs of rodent species to search for shared differences in gene expression and amino acid sequence associated with adaptation to deserts. We conducted phylogenetically independent comparisons between a desert specialist and a non-desert relative in three families representing ∼70 million years of evolution. Overall, patterns of gene expression faithfully recapitulated the phylogeny of these six taxa providing a strong evolutionary signal in levels of mRNA abundance. We also found that 8.6% of all genes showed shared patterns of expression divergence between desert and non-desert taxa, much of which likely reflects convergent evolution, and representing more than expected by chance under a model of independent gene evolution. In addition to these shared changes, we observed many species-pair-specific changes in gene expression indicating that instances of adaptation to deserts include a combination of unique and shared changes. Patterns of protein evolution revealed a small number of genes showing evidence of positive selection, the majority of which did not show shared changes in gene expression. Overall, our results suggest that convergent changes in gene regulation play an important role in the complex trait of desert adaptation in rodents.

Genomic and transcriptomic analyses reveal genetic adaptation to cold conditions in the chickens

Under the pressure of natural and artificial selection, domestic animals, including chickens, have evolved unique mechanisms of genetic adaptations such as high-altitude adaptation, hot and arid climate adaptation, and desert adaptation. Here, we investigated the genetic basis of cold tolerance in chicken by integrating whole-genome and transcriptome sequencing technologies. Genome-wide comparative analyses of 118 chickens living in different latitudes showed 46 genes and several pathways that may be involved in cold adaptation. The results of the functional enrichment analysis of differentially expressed genes proved the important role of metabolic pathways and immune-related pathways in cold tolerance in chickens. The subsequent integration of whole genome and transcriptome sequencing technology further identified six genes — dnah5 (dynein axonemal heavy chain 5), ptgs2 (prostaglandin-endoperoxide synthase 2), inhba (inhibin beta A subunit), irx2 (iroquois homeobox 2), ensgalg00000054917, and ensgalg00000046652 — requiring more detailed studies. In addition, we also discovered different allele frequency distributions of five SNPs (single nucleotide polymorphisms) within ptgs2 and nine SNPs within dnah5 in chickens in different latitudes, suggesting strong selective pressure of these two genes in chickens. We provide a novel insight into the genetic adaptation in chickens to cold environments, and provide a reference for evaluating and developing adaptive chicken breeds in cold environments.

Landscape of Global Gene Expression Reveals Distinctive Tissue Characteristics in Bactrian Camels (Camelus bactrianus)

Bactrian camels (Camelus bactrianus) are highly adapted to the desert and semi-desert environments of Asia and developed unique physiological adaptations to cold, heat, drought, and nutrient-poor conditions. These animals are an ideal model for studying desert adaptation. However, the transcriptome of different Bactrian camel tissues has not been profiled. This study performed a comprehensive transcriptome analysis of nine fetal and adult tissues. A total of 20,417 coding genes were identified, and 2.4 billion reads were generated. Gene expression and functional analyses revealed that approximately 50% of the identified genes were ubiquitously expressed, and one-third were tissue-elevated genes, which were enriched in pathways related to the biological functions of the corresponding tissue. Weighted gene co-expressed network analysis (WGCNA) identified four modules—fat metabolism, water balance, immunity, and digestion—and several hub genes, including APOA1, TMEM174, CXCL12, and MYL9. The analysis of differentially expressed genes (DEGs) between fetal and adult tissues revealed that downregulated genes were enriched in tissue development, whereas upregulated genes were enriched in biological function in adult camels. DEGs in the hump were enriched in immune-related pathways, suggesting that this tissue is involved in immunity. This study is the first to generate a transcriptome atlas of major tissues in Bactrian camels and explores the genes potentially involved in the adaptation to desert environments.

Chromosome-level genome assembly of a xerophytic plant, Haloxylon ammodendron.

Haloxylon ammodendron is a xerophytic perennial shrub or small tree that has a high ecological value in anti-desertification due to its high tolerance to drought and salt stress. Here, we report a high-quality, chromosome-level genome assembly of H. ammodendron by integrating PacBio’s high-fidelity sequencing and Hi-C technology. The assembled genome size was 685.4 Mb, of which 99.6% was assigned to nine pseudochromosomes with a contig N50 value of 23.6 Mb. Evolutionary analysis showed that both the recent substantial amplification of long terminal repeat retrotransposons and tandem gene duplication may have contributed to its genome size expansion and arid adaptation. An ample amount of low-GC genes was closely related to functions that may contribute to the desert adaptation of H. ammodendron. Gene family clustering together with gene expression analysis identified differentially expressed genes that may play important roles in the direct response of H. ammodendron to water-deficit stress. We also identified several genes possibly related to the degraded scaly leaves and well-developed root system of H. ammodendron. The reference-level genome assembly presented here will provide a valuable genomic resource for studying the genome evolution of xerophytic plants, as well as for further genetic breeding studies of H. ammodendron.

Rapidly evolving genetic features for desert adaptations in Stipagrostis pennata

BackgroundStipagrostis pennata is distributed in the mobile and semi-mobile sand dunes which can adapt well to extreme environments such as drought and high temperature. It is a pioneer plant species with potential for stabilizing sand dunes and ecological restoration. It can settle on moving sand dunes earlier than other desert plants. It can effectively improve the stability of sand dunes and help more plants settle down and increase plant diversity. However, despite its important ecological value, the genetic resources available for this species are limited.ResultsWe used single-molecule real-time sequencing technology to obtain the complete full-length transcriptome of Stipagrostis pennata, including 90,204 unigenes with an average length of 2624 bp. In addition, the 5436 transcription factors identified in these unigenes are rich in stress resistance genes, such as MYB-related, C3H, bHLH, GRAS and HSF, etc., which may play a role in adapting to desert drought and strong wind stress. Intron retention events are abundant alternative splicing events. Stipagrostis pennata has experienced stronger positive selection, accelerating the fixation of advantageous variants. Thirty-eight genes, such as CPP/TSO1-like gene, have evolved rapidly and may play a role in material transportation, flowering and seed formation.ConclusionsThe present study captures the complete full-length transcriptome of Stipagrostis pennata and reveals its rapid evolution. The desert adaptation in Stipagrostis pennata is reflected in the regulation of gene expression and the adaptability of gene function. Our findings provide a wealth of knowledge for the evolutionary adaptability of desert grass species.

Genome evolution of the psammophyte Pugionium for desert adaptation and further speciation

Deserts exert strong selection pressures on plants, but the underlying genomic drivers of ecological adaptation and subsequent speciation remain largely unknown. Here, we generated de novo genome assemblies and conducted population genomic analyses of the psammophytic genus Pugionium (Brassicaceae). Our results indicated that this bispecific genus had undergone an allopolyploid event, and the two parental genomes were derived from two ancestral lineages with different chromosome numbers and structures. The postpolyploid expansion of gene families related to abiotic stress responses and lignin biosynthesis facilitated environmental adaptations of the genus to desert habitats. Population genomic analyses of both species further revealed their recent divergence with continuous gene flow, and the most divergent regions were found to be centered on three highly structurally reshuffled chromosomes. Genes under selection in these regions, which were mainly located in one of the two subgenomes, contributed greatly to the interspecific divergence in microhabitat adaptation.

Chromosome-level genome assembly of Welwitschia mirabilis, a unique Namib Desert species.

Welwitschia mirabilis, which is endemic to the Namib Desert, is the only living species within the family Welwitschiaceae. This species has an extremely long lifespan of up to 2,000years and bears a single pair of opposite leaves that persist whilst alive. However, the underlying genetic mechanisms and evolution of the species remain poorly elucidated. Here, we report on a chromosome-level genome assembly for W.mirabilis, with a 6.30-Gb genome sequence and contig N50 of 27.50Mb. In total, 39,019 protein-coding genes were predicted from the genome. Two brassinosteroid-related genes (BRI1 and CYCD3), key regulators of cell division and elongation, were strongly selected in W.mirabilis and may contribute to their long ever-growing leaves. Furthermore, 29gene families in the mitogen-activated protein kinase signalling pathway showed significant expansion, which may contribute to the desert adaptations of the plant. Three positively selected genes (EHMT1, EIF4E, SOD2) may be involved in the mechanisms leading to long lifespan. Based on molecular clock dating and fossil calibrations, the divergence time of W.mirabilis and Gnetum montanum was estimated at ~123.5million years ago. Reconstruction of population dynamics from genome data coincided well with the aridification of the Namib Desert. The genome sequence detailed in the current study provides insight into the evolution of W.mirabilis and should be an important resource for further study on gnetophyte and gymnosperm evolution.

Genome Scan for Variable Genes Involved in Environmental Adaptations of Nubian Ibex

The Nubian ibex (Capra nubiana) is a wild goat species that inhabits the Sahara and Arabian deserts and is adapted to extreme ambient temperatures, intense solar radiation, and scarcity of food and water resources. To investigate desert adaptation, we explored the possible role of copy number variations (CNVs) in the evolution of Capra species with a specific focus on the environment of Capra nubiana. CNVs are structural genomic variations that have been implicated in phenotypic differences between species and could play a role in species adaptation. CNVs were inferred from Capra nubiana sequence data relative to the domestic goat reference genome using read-depth approach. We identified 191 CNVs overlapping with protein-coding genes mainly involved in biological processes such as innate immune response, xenobiotic metabolisms, and energy metabolisms. We found copy number variable genes involved in defense response to viral infections (Cluster of Differentiation 48, UL16 binding protein 3, Natural Killer Group 2D ligand 1-like, and Interferon-induced transmembrane protein 3), possibly suggesting their roles in Nubian ibex adaptations to viral infections. Additionally, we found copy number variable xenobiotic metabolism genes (carboxylesterase 1, Cytochrome P450 2D6, Glutathione S-transferase Mu 4, and UDP Glucuronosyltransferase-2B7), which are probably an adaptation of Nubian ibex to desert diets that are rich in plant secondary metabolites. Collectively, this study's results advance our understanding of CNVs and their possible roles in the adaptation of Nubian ibex to its environment. The copy number variable genes identified in Nubian ibex could be considered as subjects for further functional characterizations.

Integrating genetics, physiology and morphology to study desert adaptation in a lizard species.

Integration of multiple approaches is key to understand the evolutionary processes of local adaptation and speciation. Reptiles have successfully colonized desert environments, that is, extreme and arid conditions that constitute a strong selective pressure on organisms. Here, we studied genomic, physiological and morphological variations of the lizard Liolaemus fuscus to detect adaptations to the Atacama Desert. By comparing populations of L. fuscus inhabiting the Atacama Desert with populations from the Mediterranean forests from central Chile, we aimed at characterizing features related to desert adaptation. We combined ddRAD sequencing with physiological (evaporative water loss, metabolic rate and selected temperature) and morphological (linear and geometric morphometrics) measurements. We integrated the genomic and phenotypic data using redundancy analyses. Results showed strong genetic divergence, along with a high number of fixed loci between desert and forest populations. Analyses detected 110 fixed and 30 outlier loci located within genes, from which 43 were in coding regions, and 12 presented non-synonymous mutations. The candidate genes were associated with cellular membrane and development. Desert lizards presented lower evaporative water loss than those from the forest. Morphological data showed that desert lizards had smaller body size, different allometry, larger eyeballs and more dorsoventrally compressed heads. Our results suggest incipient speciation between desert and forest populations. The adaptive signal must be cautiously interpreted since genetic drift could also contribute to the divergence pattern. Nonetheless, we propose water and resource availability, and changes in habitat structure, as the most relevant challenges for desert reptiles. This study provides insights of the mechanisms that allow speciation as well as desert adaptation in reptiles at multiple levels, and highlights the benefit of integrating independent evidence.

Limited Evidence for Parallel Evolution Among Desert-Adapted Peromyscus Deer Mice.

Warming climate and increasing desertification urge the identification of genes involved in heat and dehydration tolerance to better inform and target biodiversity conservation efforts. Comparisons among extant desert-adapted species can highlight parallel or convergent patterns of genome evolution through the identification of shared signatures of selection. We generate a chromosome-level genome assembly for the canyon mouse (Peromyscus crinitus) and test for a signature of parallel evolution by comparing signatures of selective sweeps across population-level genomic resequencing data from another congeneric desert specialist (Peromyscus eremicus) and a widely distributed habitat generalist (Peromyscus maniculatus), that may be locally adapted to arid conditions. We identify few shared candidate loci involved in desert adaptation and do not find support for a shared pattern of parallel evolution. Instead, we hypothesize divergent molecular mechanisms of desert adaptation among deer mice, potentially tied to species-specific historical demography, which may limit or enhance adaptation. We identify a number of candidate loci experiencing selective sweeps in the P. crinitus genome that are implicated in osmoregulation (Trypsin, Prostasin) and metabolic tuning (Kallikrein, eIF2-alpha kinase GCN2, APPL1/2), which may be important for accommodating hot and dry environmental conditions.

Gene expression plasticity and desert adaptation in house mice.

Understanding how organisms adapt to new environments is a key problem in evolution, yet it remains unclear whether phenotypic plasticity generally facilitates or hinders this process. Here we studied evolved and plastic responses to water-stress in lab-born descendants of wild house mice (Mus musculus domesticus) collected from desert and non-desert environments and measured gene expression and organismal phenotypes under control and water-stressed conditions. After many generations in the lab, desert mice consumed significantly less water than mice from other localities, indicating that this difference has a genetic basis. Under water-stress, desert mice maintained more weight than non-desert mice, and exhibited differences in blood chemistry related to osmoregulatory function. Gene expression in the kidney revealed evolved differences between mice from different environments as well as plastic responses between hydrated and dehydrated mice. Desert mice showed reduced expression plasticity under water-stress compared to non-desert mice. Importantly, non-desert mice under water-stress generally showed shifts toward desert-like expression, consistent with adaptive plasticity. Finally, we identify several co-expression modules linked to phenotypes of interest. These findings provide evidence for local adaptation after a recent invasion and suggest that adaptive plasticity may have facilitated colonization of the desert environment.

Comparative and population genomics approaches reveal the basis of adaptation to deserts in a small rodent.

Organisms that live in deserts offer the opportunity to investigate how species adapt to environmental conditions that are lethal to most plants and animals. In the hot deserts of North America, high temperatures and lack of water are conspicuous challenges for organisms living there. The cactus mouse (Peromyscus eremicus) displays several adaptations to these conditions, including low metabolic rate, heat tolerance, and the ability to maintain homeostasis under extreme dehydration. To investigate the genomic basis of desert adaptation in cactus mice, we built a chromosome‐level genome assembly and resequenced 26 additional cactus mouse genomes from two locations in southern California (USA). Using these data, we integrated comparative, population, and functional genomic approaches. We identified 16 gene families exhibiting significant contractions or expansions in the cactus mouse compared to 17 other Myodontine rodent genomes, and found 232 sites across the genome associated with selective sweeps. Functional annotations of candidate gene families and selective sweeps revealed a pervasive signature of selection at genes involved in the synthesis and degradation of proteins, consistent with the evolution of cellular mechanisms to cope with protein denaturation caused by thermal and hyperosmotic stress. Other strong candidate genes included receptors for bitter taste, suggesting a dietary shift towards chemically defended desert plants and insects, and a growth factor involved in lipid metabolism, potentially involved in prevention of dehydration. Understanding how species adapted to deserts will provide an important foundation for predicting future evolutionary responses to increasing temperatures, droughts and desertification in the cactus mouse and other species.

A phylogenetic test of adaptation to deserts and aridity in skull and dental morphology across rodents

Understanding how organisms adapt to aridity is a central theme in traditional desert ecology research. However, many of the pioneering studies were conducted before detailed phylogenies were available to provide evolutionary context and before the accumulation of accurate bioclimatic and species distribution data to provide geographic and environmental context. We tested the desert-adaptive value of changes in skull and dental morphology in rodents after phylogenetic correction. In addition, we estimated that across the evolutionary history of more than 2,400 rodent species, transitions between mesic and desert habitats have been very frequent, with a directional bias toward the mesic-to-desert transition. This suggested that derived desert specialization is an “evolutionary deadend” that limits further evolution. After correcting for the strong phylogenetic signal, we still find a significant and strong correlation between habitat aridity and specializations associated with auditory sensitivity (auditory bulla inflation) and respiratory water retention (nasal passage elongation) but not in characters associated with dietary specialization (lower incisor shape). No other significant associations were found between habitat or aridity and any other cranial, jaw, or dental traits. Bullar hypertrophy is among the strongest patterns of convergent cranial desert adaptation in rodents and indicates that adaptation plays a similar role in shaping the evolution of this structure in different desert rodent clades.