Color Pattern Variation Research Articles

A Spatiotemporal Assessment of Dietary Partitioning between Color Morphs of a Terrestrial Salamander

When color pattern variation is co-adapted with other biological features, distinct color morphs may be separated along divergent niche axes, and data on niche partitioning can provide insight into how distinct color morphs are maintained over spatiotemporal scales. The Eastern Red-backed Salamander, Plethodon cinereus, contains two common color morphs (striped and unstriped) that differ along several trait axes in addition to coloration. Previous studies from a single population have suggested that dietary composition represents an important axis of ecological differentiation between morphs of P. cinereus. To determine if morphs partition prey resources over space and time, we collected stomach contents from morphs across six populations that ranged in color morph frequency from 100% striped to >99% unstriped, and sampled each population in a spring and a fall season. From each population and season, we also sampled leaf litter invertebrates to quantify morph differences in prey selection. Based on previous studies, we predicted striped morph diet would consist of higher quality prey in polymorphic populations, whereas in monomorphic populations, we predicted both morphs would have a more variable diet due to ecological release from intermorph competition. In the two polymorphic populations we examined, one showed no evidence of diet differences and the other mirrored differences reported from previous studies. There was no change in dietary breadth between polymorphic and monomorphic populations, and thus no signature of dietary release. Our results show there is a high degree of overlap between dietary and leaf litter invertebrates, suggesting both morphs of P. cinereus are generalist predators. Finally, we found dietary composition varied across seasons and populations, which demonstrates the importance of examining morph traits over spatial and temporal scales.

The Amazon river is a suture zone for a polyphyletic group of co‐mimetic heliconiine butterflies

The Amazon basin contains few obvious geographic barriers, yet it is the most biodiverse region on Earth. One hypothesis to explain its diversity is that the very large rivers promote allopatric divergence. Consistent with this, maps of heliconiine butterflies made from museum specimens show high subspecies richness close to the Amazon river, suggesting that it may produce or maintain intra‐specific phenotypic variability. However, museum data are subject to strong spatial biases (the ‘Wallacean shortfall' of distribution data), raising the possibility that this pattern is a sampling artefact. To test this, we systematically collected along a ~900 km north–south transect running through central Amazonia. We found a significant association between phenotypic diversity and major rivers, with distance from the Amazon river explaining 61% of the variance in the mean polymorphism of 25 species. This association is partly because many species exhibit different phenotypes on either side of the river. Nonetheless, we also find sites with high polymorphism close to the river, indicating continual cross‐river dispersal. Our results strongly suggest the presence of a suture zone (a region where multiple species have hybrid zones) near the city of Manaus. However, the effect of the river on spatial patterns of intra‐specific phenotypic diversity depends on a species' mimetic phenotype. Rather than being absolute barriers, our results support the idea that rivers can act as partial barriers that trap moving hybrid zones, resulting in a suture zone. As such, the wide Amazonian rivers help generate and maintain colour pattern diversity, but to date there is no evidence that they lead to speciation in our study group.

Deep Convergence, Shared Ancestry, and Evolutionary Novelty in the Genetic Architecture of Heliconius Mimicry.

Convergent evolution can occur through different genetic mechanisms in different species. It is now clear that convergence at the genetic level is also widespread, and can be caused by either (i) parallel genetic evolution, where independently evolved convergent mutations arise in different populations or species, or (ii) collateral evolution in which shared ancestry results from either ancestral polymorphism or introgression among taxa. The adaptive radiation of Heliconius butterflies shows color pattern variation within species, as well as mimetic convergence between species. Using comparisons from across multiple hybrid zones, we use signals of shared ancestry to identify and refine multiple putative regulatory elements in Heliconius melpomene and its comimics, Heliconius elevatus and Heliconius besckei, around three known major color patterning genes: optix, WntA, and cortex While we find that convergence between H. melpomene and H. elevatus is caused by a complex history of collateral evolution via introgression in the Amazon, convergence between these species in the Guianas appears to have evolved independently. Thus, we find adaptive convergent genetic evolution to be a key driver of regulatory changes that lead to rapid phenotypic changes. Furthermore, we uncover evidence of parallel genetic evolution at some loci around optix and WntA in H. melpomene and its distant comimic Heliconius erato Ultimately, we show that all three of convergence, conservation, and novelty underlie the modular architecture of Heliconius color pattern mimicry.

High-resolution characterization of male ornamentation and re-evaluation of sex linkage in guppies.

Coloration plays a key role in the ecology of many species, influencing how an organism interacts with its environment, other species and conspecifics. Guppies are sexually dimorphic, with males displaying sexually selected coloration resulting from female preference. Previous work has suggested that much of guppy colour pattern variation is Y-linked. However, it remains unclear how many individual colour patterns are Y-linked in natural populations as much of the previous work has focused on phenotypes either not found in the wild, or aggregate measures such as total colour area. Moreover, ornaments have traditionally been identified and delineated by hand, and computational methods now make it possible to extract pixels and identify ornaments with automated methods, reducing the potential for human bias. Here we developed a pipeline for semi-automated ornament identification and high-resolution image analysis of male guppy colour patterns and applied it to a multigenerational pedigree. Our results show that loci controlling the presence or the absence of individual male ornaments in our population are not predominantly Y-linked. However, we find that ornaments of similar colour are not independent of each other, and modifier loci that affect whole animal coloration appear to be at least partially Y-linked. Considering these results, Y-linkage of individual ornaments may not be important in driving colour changes in natural populations of guppies, or in expansions of the non-recombining Y region, while Y-linked modifier loci that affect aggregate traits may well play an important role.

A new species of the porcelain crab genus Petrolisthes from the Pacific coast of Panama, with taxonomic notes on closely related species (Decapoda: Anomura: Porcellanidae).

A new species of porcelain crab, Petrolisthes lazarus sp. nov., is described from the Pacific coast of Panama, based on two female specimens. The new species is morphologically most similar to another eastern Pacific species, P. crenulatus Lockington, 1878, especially in the general configuration of the carapace, chelipeds and ambulatory legs. However, P. lazarus sp. nov. can be separated from P. crenulatus by the differences in the proportions of the carapace, the shape of the frontal region of the carapace, and the setation pattern of the cheliped. In addition, P. lazarus sp. nov. and P. crenulatus appear to be allopatric, the latter species presently being known only from Mexico. Among other eastern Pacific taxa, P. lazarus sp. nov. may have some affinities with P. ortmanni Nobili, 1901 and P. lewisi (Glassell, 1936), from which it can be easily distinguished by the chelipeds lacking a dense field of setae on the outer (lateral) surface of the palm and with different proportions of some articles, especially the carpus. The four species can also be distinguished from each other by their diagnostic, although somewhat variable colour patterns.

Spatial and temporal dynamics of exuberant colour polymorphism in the southern cricket frog

ABSTRACT Intraspecific variation in colour pattern is widespread across multitudinous amphibian species. In some species, many distinct colour patterns are maintained within populations, a phenomenon referred to as exuberant colour polymorphism. The underlying causes of exuberant colour pattern polymorphism are poorly understood but are likely explained by selection, rather than neutral processes like genetic drift. Nevertheless, empirical data are needed to understand the selective drivers of this phenomenon, but such data are currently lacking for most polymorphic species. We studied frequency, spatial, and linkage dynamics of colour pattern across nine populations of the southern cricket frog (Acris gryllus) from southeastern Georgia, USA. Using 233 individuals, we combined direct field observations with examination of natural history specimens to look at colour pattern characteristics as they relate to space, time, and sex over a 30-year time frame. We found evidence of spatial and temporal variation in colour pattern across populations. We also discovered associations among colour pattern traits and between two colour pattern traits and sex. Our results suggest that the exuberant colour polymorphism of A. gryllus may involve correlations between traits and be caused by spatial and/or temporal variation in selection. However, similar studies in other species are necessary to allow us to discriminate among different drivers of colour pattern in exuberantly polymorphic frogs. Collectively, such systems offer important opportunities for understanding the evolution of colour and phenotypic diversity.

Divergence, gene flow, and the origin of leapfrog geographic distributions: The history of colour pattern variation in Phyllobates poison-dart frogs.

The geographic distribution of phenotypic variation among closely related populations is a valuable source of information about the evolutionary processes that generate and maintain biodiversity. Leapfrog distributions, in which phenotypically similar populations are disjunctly distributed and separated by one or more phenotypically distinct populations, represent geographic replicates for the existence of a phenotype, and are therefore especially informative. These geographic patterns have mostly been studied from phylogenetic perspectives to understand how common ancestry and divergent evolution drive their formation. Other processes, such as gene flow between populations, have not received as much attention. Here, we investigate the roles of divergence and gene flow between populations in the origin and maintenance of a leapfrog distribution in Phyllobates poison frogs. We found evidence for high levels of gene flow between neighbouring populations but not over long distances, indicating that gene flow between populations exhibiting the central phenotype may have a homogenizing effect that maintains their similarity, and that introgression between 'leapfroging' taxa has not played a prominent role as a driver of phenotypic diversity in Phyllobates. Although phylogenetic analyses suggest that the leapfrog distribution was formed through independent evolution of the peripheral (i.e. leapfrogging) populations, the elevated levels of gene flow between geographically close populations poise alternative scenarios, such as the history of phenotypic change becoming decoupled from genome-averaged patterns of divergence, which we cannot rule out. These results highlight the importance of incorporating gene flow between populations into the study of geographic variation in phenotypes, both as a driver of phenotypic diversity and as a confounding factor of phylogeographic inferences.

Evolution of Pelage Luminance in Squirrels (Sciuridae)

Pelage luminance has been found in many mammalian systems to follow patterns predicted by Gloger’s rule where darker colored animals are associated with environments that are warmer and more moist. Sciurids have one of the greatest diversities of color patterns and hues among mammalian families. We have used comparative methods to investigate whether the luminance of dorsal pelage in 137 species across Sciuridae conform to prediction of Gloger’s rule and other background matching expectations. We found using phylogenetic multiple regression, as well as univariate regression, that Sciurids generally conform to expectations of Gloger’s rule. Darker species are associated with environments with higher primary productivity, higher temperature, higher humidity, and lower solar radiation. Moreover, in support of the predictions of background matching, darker squirrel species were associated with environments with greater soil carbon content and higher fire frequency. Our macroevolutionary study sheds some light on selective pressures that are driving the evolution of coloration in Sciurids, but more comparative research is needed to fully understand other selective pressures that have led to the wide diversity of color patterns and hues.

Adding stars to the Chromodoris (Nudibranchia, Chromodorididae) galaxy with the description of four new species.

This paper describes four new Chromodoris species: Chromodoris balat sp. nov., Chromodoris baqe sp. nov., Chromodoris kalawakan sp. nov., and Chromodoris quagga sp. nov. We were able to distinguish 44 species level lineages within Chromodoris, expanding the Indo-Pacific species from 39 species. The phylogeny presented here provides slightly greater resolution of species relationships than do previous studies of this genus. Layton et al. (2018), determined that variable color patterns made it difficult to differentiate in living specimens of distinct species, but we provide some additional stable color characters that potentially help resolve this issue, as well as additional internal features that are useful in species delimitation and correlate with the molecular phylogenetic analysis.

Perfect mimicry between Heliconius butterflies is constrained by genetics and development.

Müllerian mimicry strongly exemplifies the power of natural selection. However, the exact measure of such adaptive phenotypic convergence and the possible causes of its imperfection often remain unidentified. Here, we first quantify wing colour pattern differences in the forewing region of 14 co-mimetic colour pattern morphs of the butterfly species Heliconius erato and Heliconius melpomene and measure the extent to which mimicking colour pattern morphs are not perfectly identical. Next, using gene-editing CRISPR/Cas9 KO experiments of the gene WntA, which has been mapped to colour pattern diversity in these butterflies, we explore the exact areas of the wings in which WntA affects colour pattern formation differently in H. erato and H. melpomene. We find that, while the relative size of the forewing pattern is generally nearly identical between co-mimics, the CRISPR/Cas9 KO results highlight divergent boundaries in the wing that prevent the co-mimics from achieving perfect mimicry. We suggest that this mismatch may be explained by divergence in the gene regulatory network that defines wing colour patterning in both species, thus constraining morphological evolution even between closely related species.

Macroevolution of Flower Color Patterning: Biased Transition Rates and Correlated Evolution with Flower Size.

Floral pigmentation patterns can both mediate plant-pollinator interactions and modify the abiotic environment of reproductive structures. To date, there have been no inquiries into the rate and directionality of macroevolutionary transitions between patterned and non-patterned petals despite their ecological importance and ubiquity across angiosperms. Petals in the Potentilleae tribe (Rosaceae) display color patterns in the ultraviolet (UV) and human-visible spectrum, or can be uniform in color (i.e., patternless). Using a phylogeny of Potentilleae, I test whether evolutionary transition rates between patterned and non-patterned petals are biased in either direction. I then examine whether UV and human-visible floral patterns are phylogenetically correlated and test the prediction that color patterns will evolve in concert with larger flowers if they function as guides to orient pollinators to floral rewards. I found that transition rates were biased toward petals that were uniform in color. Transition rates from patterned to uniformly colored petals were two and six times higher than the reverse for UV and human-visible pattern, respectively. The presence of UV and human-visible pattern evolved independently from one another. However, the evolution of human-visible pattern was associated with the evolution of larger flowers but the evolution of UV pattern was correlated with the evolution of smaller flowers. I posit that the transition bias toward non-patterned flowers may reflect developmental constraints on spatial regulation of pigments required to produce floral color patterning. The correlated evolution of larger flowers and human-visible pigmentation patterns support the hypothesis that nectar or pollen guides are more likely to evolve in larger-flowered species. This work provides insight into how transition rate bias and trait correlations can shape phylogenetic patterns of floral color pattern diversity.

Ontogenetic changes in the ventral colouration of post metamorphic Elachistocleis haroi Pereyra, Akmentins, Laufer, Vaira, 2013 (Anura: Microhylidae)

Abstract Ontogenetic colour change (OCC) is defined as the progressive and non-reversible process of changes in colouration of organisms associated with their development. Among the many vertebrate groups, amphibians are particularly impressive for their strikingly wide variety of colours, colour patterns, and signals, whose evolutionary and ecological significance have been poorly studied. Elachistocleis comprises 18 species currently separated into two main groups based on their ventral colour pattern: one immaculate and the other with specks and/or colour patches. Elachistocleis haroi is a small-sized species within the immaculate venter group, distributed in the Yungas and Dry Chaco ecoregions from which little information is known. In a comprehensive sampling of post-metamorphic individuals of E. haroi at different stages of development we identified a significant variation in ventral colour pattern, which could denote a progressive filling of yellow colour according to an ontogenetic pattern. To test this hypothesis, we analysed 39 post-metamorphic individuals of E. haroi at different stages of development with imaging procedures. We found that yellow spots and their intensity are significantly related to snout-vent length, as major expansion of colour on the sides, gular region and male chest, as almost no development on the belly. We briefly discuss our findings in relation to sexual display and predation avoidance. To our knowledge, this is the first analysis of post-metamorphic OCC in ventral colouration in the genus Elachistocleis.

Urban moth communities suggest that life in the city favours thermophilic multi-dimensional generalists.

Biodiversity is challenged worldwide by exploitation, global warming, changes in land use and increasing urbanization. It is hypothesized that communities in urban areas should consist primarily of generalist species with broad niches that are able to cope with novel, variable, fragmented, warmer and unpredictable environments shaped by human pressures. We surveyed moth communities in three cities in northern Europe and compared them with neighbouring moth assemblages constituting species pools of potential colonizers. We found that urban moth communities consisted of multi-dimensional generalist species that had larger distribution ranges, more variable colour patterns, longer reproductive seasons, broader diets, were more likely to overwinter as an egg, more thermophilic, and occupied more habitat types compared with moth communities in surrounding areas. When body size was analysed separately, results indicated that city occupancy was associated with larger size, but this effect disappeared when body size was analysed together with the other traits. Our findings indicate that urbanization imposes a spatial filtering process in favour of thermophilic species characterized by high intraspecific diversity and multi-dimensional generalist lifestyles over specialized species with narrow niches.

Transcription factors underlying wing margin color patterns and pupal cuticle markings in butterflies

BackgroundThe diversity of butterfly color patterns can be attributed to a relatively small number of pattern elements that are homologous across Lepidoptera. Although genes involved in patterning some of these elements have been identified, the development of several major elements remains poorly understood. To identify genes underlying wing pupal cuticle markings and wing margin color patterns, we examined expression of the candidate transcription factors Engrailed/Invected (En/Inv), Distal-less (Dll), Cubitus interruptus (Ci), and Spalt in two nymphalids: Junonia coenia and Bicyclus anynana.ResultsWe found that En/Inv, Dll, and Ci mark domains on the J. coenia last-instar forewing disc that closely correspond to the position and shape of pupal cuticle markings. We also found that Spalt demarcates wing margin color patterns in both J. coenia and B. anynana, and that CRISPR/Cas9 deletions in the spalt gene result in reduction and loss of wing margin color patterns in J. coenia. These data demonstrate a role for spalt in promoting wing margin color patterning, in addition to its previously described role in eyespot patterning.ConclusionOur observations support the model that a core set of regulatory genes are redeployed multiple times, and in multiple roles, during butterfly wing pattern development. Of these genes, spalt is of special interest as it plays a dual role in both eyespot and margin color pattern development.

Hybridization and transgressive exploration of colour pattern and wing morphology in Heliconius butterflies.

Hybridization can generate novel phenotypes distinct from those of parental lineages, a phenomenon known as transgressive trait variation. Transgressive phenotypes might negatively or positively affect hybrid fitness, and increase available variation. Closely related species of Heliconius butterflies regularly produce hybrids in nature, and hybridization is thought to play a role in the diversification of novel wing colour patterns despite strong stabilizing selection due to interspecific mimicry. Here, we studied wing phenotypes in first- and second-generation hybrids produced by controlled crosses between either two co-mimetic species of Heliconius or between two nonmimetic species. We quantified wing size, shape and colour pattern variation and asked whether hybrids displayed transgressive wing phenotypes. Discrete traits underlain by major-effect loci, such as the presence or absence of colour patches, generate novel phenotypes. For quantitative traits, such as wing shape or subtle colour pattern characters, hybrids only exceed the parental range in specific dimensions of the morphological space. Overall, our study addresses some of the challenges in defining and measuring phenotypic transgression for multivariate traits and our data suggest that the extent to which transgressive trait variation in hybrids contributes to phenotypic diversity depends on the complexity and the genetic architecture of the traits.

Functional colour genes and signals of selection in colour-polymorphic salamanders.

Coloration has been associated with multiple biologically relevant traits that drive adaptation and diversification in many taxa. However, despite the great diversity of colour patterns present in amphibians the underlying molecular basis is largely unknown. Here, we use insight from a highly colour-variable lineage of the European fire salamander (Salamandra salamandra bernardezi) to identify functional associations with striking variation in colour morph and pattern. The three focal colour morphs-ancestral black-yellow striped, fully yellow and fully brown-differed in pattern, visible coloration and cellular composition. From population genomic analyses of up to 4,702 loci, we found no correlations of neutral population genetic structure with colour morph. However, we identified 21 loci with genotype-phenotype associations, several of which relate to known colour genes. Furthermore, we inferred response to selection at up to 142 loci between the colour morphs, again including several that relate to coloration genes. By transcriptomic analysis across all different combinations, we found 196 differentially expressed genes between yellow, brown and black skin, 63 of which are candidate genes involved in animal coloration. The concordance across different statistical approaches and 'omic data sets provide several lines of evidence for loci linked to functional differences between colour morphs, including TYR, CAMK1 and PMEL. We found little association between colour morph and the metabolomic profile of its toxic compounds from the skin secretions. Our research suggests that current ecological and evolutionary hypotheses for the origins and maintenance of these striking colour morphs may need to be revisited.

Contrasting genomic and phenotypic outcomes of hybridization between pairs of mimetic butterfly taxa across a suture zone.

Hybrid zones, whereby divergent lineages come into contact and eventually hybridize, can provide insights on the mechanisms involved in population differentiation and reproductive isolation, and ultimately speciation. Suture zones offer the opportunity to compare these processes across multiple species. In this paper we use reduced-complexity genomic data to compare the genetic and phenotypic structure and hybridization patterns of two mimetic butterfly species, Ithomia salapia and Oleria onega (Nymphalidae: Ithomiini), each consisting of a pair of lineages differentiated for their wing colour pattern and that come into contact in the Andean foothills of Peru. Despite similarities in their life history, we highlight major differences, both at the genomic and phenotypic level, between the two species. These differences include the presence of hybrids, variations in wing phenotype, and genomic patterns of introgression and differentiation. In I.salapia, the two lineages appear to hybridize only rarely, whereas in O.onega the hybrids are not only more common, but also genetically and phenotypically more variable. We also detected loci statistically associated with wing colour pattern variation, but in both species these loci were not over-represented among the candidate barrier loci, suggesting that traits other than wing colour pattern may be important for reproductive isolation. Our results contrast with the genomic patterns observed between hybridizing lineages in the mimetic Heliconius butterflies, and call for a broader investigation into the genomics of speciation in Ithomiini - the largest radiation of mimetic butterflies.

Inconspicuous genetic and morphological patterns challenge the taxonomic status of endemic species Bodianus insularis (Labridae)

Bodianus constitutes a marine fish group of particular interest due to its trophic interactions with reef environments. Knowledge of phylogenetic relationships of this genus have been improved by shared advances of classical and genetic approaches. However, some of its species have yet to be compared regarding its genetic patterns. Morphological variations in body shape and color patterns during the course of ontogenetic development, in addition to recurring sex reversal in species of this genus, make its taxonomy challenging. In the Atlantic Ocean five species have been described: Bodianus scrofa and Bodianus speciosus (Eastern Atlantic), Bodianus rufus (Western Atlantic), Bodianus pulchellus (Western Atlantic and Eastern Atlantic Islands) and Bodianus insularis (Mid-Atlantic islands). The phylogenetic relationships of the last species could not be resolved using genetic analyses. With a very restrict geographic distribution, B. insularis deserves high attention regarding its taxonomic and evolutionary aspects. Here, we use mitochondrial DNA sequences (COI and 16S rRNA) and nuclear analyses (rhodopsin) to assess the genetic divergence of B. insularis in relation to other Atlantic species and clarify its phylogenetic relationships. Bodianus pulchellus and B. insularis were not genetically distinct and are grouped with B. rufus, forming a sister clade of B. speciosus, while B. scrofa is more related to Pacific species. DNA-based and morphological traits are very similar between B. insularis and B. pulchellus, whose most obvious difference is their color pattern. The actual reinterpretation of ecological and biogeographic contexts allows to suggest that B. insularis is most likely a synonym of B. pulchellus, constituting a population enclave of this species.

A Survey of the Floral Volatile Profiles of Daylily Species and Hybrids

For the past century, daylily (Hemerocallis) hybridization focused almost exclusively on enhancing the diversity of flower forms, colors, and color patterns. This focus on the visual characteristics of daylilies resulted in the development of thousands of hybrids that come in a fantastic array of colors and color patterns. However, these modern daylilies exhibit little to none of the floral fragrance possessed by some of the daylily progenitor species. Because little work has been done on daylily floral volatiles, the objective of this research was to evaluate the floral volatile profiles of a variety of daylily species and hybrids and assess the state of fragrance among modern hybrids. Using a field collection system and gas chromatography–mass spectrometry (GS-MS), this study evaluated the fragrance profiles of 147 daylily genotypes. Eighteen volatile organic compounds, primarily terpenoids, were identified and their variations among the genotypes were investigated. Results suggest that fragrance could be a trait pursued in a breeding program to enhance the sensory phenotypes of new daylily cultivars.

Genetic differentiation and overexploitation history of the critically endangered Lehmann’s Poison Frog: Oophaga lehmanni

Species conservation with fragmented and endangered populations must be based on a prior and thorough knowledge of the structure and population dynamics. Oophaga lehmanni is a dendrobatid species endemic of Colombia and is restricted to its type locality. This species has a fragmented distribution and is considered as critically endangered mainly due to habitat destruction and overexploitation. Oophaga lehmanni exhibits phenotypic variation in the dorsal color pattern (red and yellow morphs). We reconstructed the overexploitation history that this species has faced in the last 40 years. In addition, we collected genetic and morphological data for the first time in natural populations to describe genetic diversity between and within populations, and to evaluate morphological and genetic differences between red and yellow morphs. Overexploitation data suggest that more than 80.000 (Min = 60.047–Max = 102.236) frogs were extracted from the field in the last four decades, probably generating the local extirpation or population decline from the type locality. Genetic data showed reduced genetic diversity. Observed heterozygosity (mean ± s.d. = 0.599 ± 0.165) is lower than expected (mean ± s.d. = 0.867 ± 0.082). We did not find differences in body size and heterozygosity between the two morphs; however, individuals analyzed were assigned to two genetic clusters, which corresponded to the O. lehmanni-yellow and O. lehmanni-red. In addition, FST (0.209) and Nei genetic distance (0.18) values indicated genetic differentiation between the two morphs; therefore, red and yellow morphs should be treated as independent management units. This information will help to define appropriate and long-term conservation units, as a useful tool to mitigate the extinction risk of this species.