Biology subjects

Salamin, N.

Publications and source records attributed to Salamin, N..

At least 19 recordsLinked to original sources

bioRxiv2026

Heterogeneous evolution of effective recombination landscapes across percomorph fishes

Recombination shapes allelic combinations across the genome and plays a central role in adaptation and speciation. While recombination rates are known to vary and evolve, how recombination landscapes diverge between populations and which processes drive this divergence remains poorly understood. Here, we use the Population Recombination Divergence Index (PRDI) to quantify divergence in effective recombination landscapes between seven pairs of populations and species from four percomorph fish lineages (cichlids, sticklebacks, killifish, and clownfish). While accounting for stochasticity and confounders of linkage disequilibrium (LD) -based recombination rate estimates, we find that recombination landscapes diverged substantially in all comparisons. This divergence generally increases with genetic differentiation and divergence time, but the amount of PRDI accumulated per unit of genetic divergence (FST), net nucleotide divergence (da), or divergence time varies markedly among population and species pairs. Comparisons across systems suggest that PRDI accumulation tends to be faster (per unit of FST, da, or divergence time) in pairs with higher inferred gene flow. We also introduce a local version of PRDI (_l) to better characterize genomic heterogeneity in effective recombination divergence. We find that higher inferred gene flow between populations tends to be associated with stronger intragenomic correlation between local FST and _l. Finally, we propose a conceptual framework for investigating how divergent selection and gene flow may interact to shape effective recombination landscape divergence. Our work highlights the heterogeneous dynamics of effective recombination landscape evolution, and motivates further theoretical and empirical efforts to integrate recombination landscape evolution into studies of adaptation and speciation.

evolutionary biology↗

bioRxiv2026

Genomic structural variation in clownfish adaptive radiation

Clownfishes are a complex of 28 species that rapidly diversified after acquiring mutualism with host sea anemones around 15 million years ago. The genomic mechanisms behind this radiation are only beginning to emerge, and structural variants (SVs) have received little attention. Previous phylogenomic studies suggested two large inversions on chromosome 18, but their distribution across the clownfish phylogeny and their potential effects have not been formally explored. SVs can disrupt gene function and regulation or alter gene dosage, and they have been shown to play a central role in adaptive evolution and diversification across many taxa. Here, we characterized SVs across the clownfish radiation using long-read PacBio sequencing, generating high-quality genome assemblies for sixteen species spanning the main lineages of the genus. We identified and catalogued nearly 130,000 SVs spanning close to 570 Mb. Short insertions and deletions were the most numerous, but large inversions accounted for most of the affected sequence. No SV was consistently associated with host specialization. We nevertheless confirmed a single inversion of approximately 17 Mb on chromosome 18, clarifying earlier reports of two separate rearrangements at this locus, and identified an additional inversion on chromosome 9 whose distribution suggests a history of hybridization. This study provides the first genome-wide characterization of structural variation in clownfishes, revealing extensive SVs despite the group's rapid and recent radiation. It also delivers new, high-quality genomic resources that will support future research into this iconic group of coral reef fishes and their adaptation to sea anemones.

genomics↗

bioRxiv2026

One species or several? Genomic evidence for deep divergence within the widely distributed anemonefish Amphiprion clarkii species complex

Defining what a species is remains one of the biggest questions in biology and directly challenges species delimitation in species complexes. The wide-spread Clarks' anemonefish Amphiprion clarkii has the widest range of all anemonefishes, from the Indian Ocean to the Central Pacific. However, the taxonomy of this species has been confused since its initial description due to its great variation in morphology and pigmentation even within the same locality. Here, we combined whole genome re-sequencing data from 166 individuals spanning most of the distribution of A. clarkii, including remote islands such as Ogasawara and Guam in the northwestern Pacific, and also including three individuals of the closely related A. tricinctus from the Marshall Islands. Our population genomic analyses reveal an unexpected level of divergence, with five deeply separated lineages corresponding to the Indian Ocean, Central Indo-Pacific, Melanesia, and the isolated population of Ogasawara. The fifth lineage comprises A. clarkii from Guam and A. tricinctus, revealing that A. tricinctus is phylogenetically nested within A. clarkii, challenging its current taxonomic status. The most divergent lineage, Melanesia, is genetically more distant from other A. clarkii lineages than several recognised species pairs within the Amphiprion genus, yet demographic modelling shows ongoing gene flow between all lineages. Together, our results highlight a hidden diversity and stress the necessity of an integrative taxonomic revision of the A. clarkii species complex.

evolutionary biology↗

bioRxiv2026

Discovery and genomic characterization of a novel wild clownfish hybrid from the Philippines

Hybridization is widespread among marine fishes and can contribute to adaptation, diversification, and the generation of novel phenotypes. In clownfishes, only two wild hybrids Amphiprion leucokranos and A. thiellei have been described, yet those naturally occurring clownfish hybrids remain rarely documented. Both hybrids involve crosses with A. sandaracinos. During field surveys in the Philippines, we identified a previously undocumented clownfish individual with an unusual phenotype resembling both A. sandaracinos and A. perideraion. To characterize its origin, we combined genomic, mitochondrial, and phenotypic comparisons to previously described clownfish hybrid systems. Genome-wide PCA and admixture analyses supported mixed ancestry between A. sandaracinos and A. perideraion. Reconstruction of the mitochondrial genome placed the individual within the A. sandaracinos mitochondrial lineage. Together, these results support a hybrid origin and suggest predominant A. sandaracinos ancestry, consistent with a backcrossed descendant rather than a first-generation hybrid. Comparisons with the previously characterized hybrid A. leucokranos further revealed similarities in genomic composition and phenotype across independently derived clownfish hybrid systems. Our findings identify a previously undocumented natural clownfish hybrid and suggest that integrating genomic and field-based approaches may reveal additional cryptic hybrid systems and improve understanding of hybridization in clownfishes.

genomics↗

bioRxiv2026

Color polymorphism in the saddleback clownfish, Amphiprion polymnus: species or complex?

Color polymorphism can facilitate local adaptation, maintain intraspecific diversity, or reflect early stages of speciation. Clownfishes (Amphiprion spp.) typically display a simple black, orange, and white pattern, but the saddleback clownfish (Amphiprion polymnus) shows striking variation in melanism and the number of vertical bars, which are thought to play a role in species recognition. In 2024, a revision on iNaturalist split A. polymnus into multiple species based solely on color pattern and geographic range. This raises the question of whether these morphs represent true species or intraspecific polymorphism, which we tested using genomic and image-based data. We sampled 97 individuals from seven populations across the species range and quantified color patterns from standardized photographs. Phenotypic and genomic analyses reveal a complex pattern of divergence. Image analysis identified three distinct phenotypic clusters, with A. polymnus, A. annamensis, and A. laticlavius each showing consistent differences in saddle shape and vertical bar extent. ADMIXTURE resolved three distinct genetic groups corresponding to the morphs. Pairwise FST (0.54-0.71) and dxy indicate extremely high differentiation between A. polymnus and A. annamensis, consistent with species-level divergence, whereas A. laticlavius shows much lower differentiation from A. polymnus (FST 0.09-0.18) and higher differentiation from A. annamensis (FST 0.64-0.66). Overall, phenotypic and genomic data show structured variation, but the status of A. laticlavius remains ambiguous. Our study reveals clear and structured divergence across the full range, yet the taxonomic interpretation of this variation remains inherently challenging. The key question remains: do these patterns reflect a single polymorphic species or a complex of closely related species?

ecology↗

bioRxiv2025

Colour and shape evolution reflect ecological specialisation in Pomacentridae

Damselfishes (Pomacentridae) display remarkable diversity in colouration and body form, yet the processes shaping this phenotypic variation remain poorly resolved. Our study aimed to characterise the evolution of these traits, evaluate their associations with ecological factors, and identify convergent patterns linked to ecological specialisation. Using image-based quantification of colour patterns, geometric morphometrics, and phylogenetic comparative methods across 343 species, we show that pomacentrid phenotypes are organised around a small number of dominant axes describing brightness, hue, contrast, body elongation, and cranial morphology. Both colour and morphology exhibit early bursts of evolutionary disparity, followed by recurrent lineage-specific radiations and widespread convergence toward similar adaptive optima across the phylogeny. Dietary ecotypes emerged as the strongest predictor of morphological diversification, whereas symbiotic and social regimes exerted the strongest effects on colour evolution. Despite these distinct ecological correlates, several colour and shape axes form partially integrated trait syndromes that evolve in concert. The pervasive convergence of colour-shape syndromes underscores deterministic components of reef-fish evolution and positions Pomacentridae as a model for understanding integrated phenotypic evolution.

evolutionary biology↗

bioRxiv2025

Phylogram instead of chronogram when assessing the neutral evolution of a trait

Comparative analyses of trait evolution aim to uncover the different evolutionary forces shaping phenotypic diversity among species. This is typically done by fitting various evolutionary models to observed trait changes along a species tree. For instance, under neutral evolution, trait values are modelled as changing randomly along the branches of the tree. In contrast, for a trait under selection, species are typically assumed to track an optimal trait value, which itself may shift along the tree. Here, rather than relying on alternative models to discriminate among evolutionary scenarios, we focus on the underlying species tree, and specifically the units in which its branch lengths are measured. Species trees are usually represented as chronograms, with branch lengths proportional to time. The rationale is that time correlates with trait changes through its connection to the number of generations. However, since the generation time of species can also vary along the phylogenetic tree, we argue that chronograms introduces biases. In contrast, phylograms with branch lengths in units of sequence divergence will account for the effect of changing generation time. In this study, we develop a method to test whether a phylogram provides a better fit than a chronogram for modelling trait evolution. We show using simulations that, for a trait evolving neutrally, the fit of a random evolution model has more support on a phylogram than on a chronogram. However, comparing models and testing different scenarios of selection using a phylogram leads to incorrect predictions. Given these results, we argue that we should use phylograms instead of chronograms when assessing the neutral evolution of a trait. Nevertheless, we support the fact that we should generally continue to use chronograms to model selection acting on a quantitative trait. Lay summaryWhen we look at different species, we can see that some traits are different between them. For example, the brain size of different primate and human lineages varies. The question we ask is: does the time that passed explain the differences in their traits? Or are genetic differences between species better at explaining the differences? The standard in evolutionary biology is to use time to explain differences in traits. However, if the trait is not under natural selection and instead is changing due to accumulating mutations, the genetic differences between species instead of time should best explain the differences in traits. As a result, we argue that using both genetic differences and species divergence time simultaneously provides a better understanding of how traits evolve across species.

evolutionary biology↗

bioRxiv2025

Environmental fluctuations trigger faster evolution and turnover

Biodiversity patterns and species communities emerge from the interactions between organisms and their environment and from processes spanning across different spatial and temporal scales. Understanding the dynamics of these interactions remains difficult as it requires mapping how microevolutionary processes scale up to macroevolutionary outcomes. Here, we use an individual-based simulation framework to model how populations respond to environmental fluctuations in a spatially explicit context. Our approach tracks evolutionary processes over thousands of generations to quantify the emerging macroevolutionary patterns under varying degrees of environmental instability modeled as shifting environmental gradients. We found that increased instability across time and space leads to elevated rates of species-level extinctions leading to homogenization of species communities, while promoting high rates of phenotypic evolution. Emerging extinction patterns indicate a selective process where species with low migration ability, narrow environmental tolerance, and limited evolvability are disproportionally affected by phases of rapid environmental change. Understanding the impact of environmental variation on organisms and species evolution remains challenging but can be explored through simulations to generate expectations and testable hypotheses about evolutionary processes across different scales.

evolutionary biology↗

bioRxiv2025

Genetic basis of the evolution of vertical bars in clownfishes

Clownfish exhibit striking color patterns, characterized primarily by the presence of zero to three vertical white bars, along with three main colors: orange, white, and black. The common ancestor of clownfish likely possessed three vertical bars, with several instances of gains and losses occurring throughout clownfish evolutionary history over the past 10 million years. However, the evolutionary genomic mechanisms underlying the gain or loss of vertical bars remain unknown. In this study, we tested whether vertical bar transitions across the clownfish phylogeny were associated with changes in non-synonymous to synonymous substitution rates (dN/dS values). Our analyses identified pigmentation-related genes that underwent changes in selective pressure, including gch2, oca2, and vps11, which are linked to melanophores, iridophores, and visual function. Additionally, pmel, a key melanogenesis gene, was found under positive selection, suggesting its role in shaping bar patterning. These results provide new insights into the genomic basis of coloration in clownfish, highlighting how selection and genetic variation influence phenotypic evolution.

evolutionary biology↗

bioRxiv2024

Widespread selection relaxation in aquatic mammals

While mammals are predominantly terrestrial, several lineages within them have independently transitioned to aquatic environments and exhibit a great variety of evolutionary changes, which ultimately lead to irreversible transitions to aquatic lifestyle. Such changes should also be detectable at the molecular level and, for instance, olfactory genes have been found to evolve under reduced selection and functionality in whales. However, the presence of a molecular signature related to aquatic transitions across other genes remains unknown as is the degree to which this affects only fully aquatic groups or also semiaquatic ones. Here, we use a Bayesian framework to investigate differences in the strength of purifying selection among terrestrial, semiaquatic and fully aquatic mammals using a set of 1000 orthologous genes, while accounting for shared ancestry, and controlling for body mass and effective population size. We found that selection relaxation linked to aquatic transitions is not only occurring in olfactory genes, but also detected across 20% of the other genes analysed here. This consistent trend of genes under weaker selection is inferred in both semi- and fully aquatic mammals although the pattern is stronger in the latter. While most of the genes analysed here likely remain functional in all mammals, their evolution is inferred to be neutral in a substantial number of aquatic species, consistently with genes that might be losing their functionality. The inferred widespread relaxation of selection among genes is consistent with a macroevolutionary scenario where secondary transitions to aquatic environment eventually become irreversible.

evolutionary biology↗

bioRxiv2024

Specialization into Host Sea Anemones Impacted Clownfish Demographic Responses to Pleistocene Sea Level Changes

Fluctuating sea levels during the Pleistocene led to habitat loss and fragmentation, impacting the evolutionary trajectories of reef fishes. Species with specialized ecological requirements or habitat preferences, like clownfishes (Amphiprioninae), may have been particularly vulnerable due to their intricate dependence on sea anemones. The diverse host specializations within this group likely contributed distinct responses to sea-level fluctuations, differentially shaping their recent evolutionary histories. Leveraging a comprehensive genomic dataset, we reveal demographic patterns and connectivity dynamics across multiple populations of ten clownfish species under different host specializations. Host-generalist species demonstrated strong resilience to habitat perturbations, while those specialized on single hosts suffered dramatic bottlenecks linked to sea-level fluctuations. Spatial analyses revealed the significant role of oceanic currents in shaping clownfish genetic diversity landscapes. Dispersal barriers were driven by environmental variables, with the Coral Triangle emerging as a hub of genetic diversity. Our results reveal how clownfish associative behavior influences their population dynamics, holding major implications for their conservation such as the need to consider their mutualism with sea anemones, particularly on host-specialists, to ensure their survival in the face of climate threats. These findings extend broader principles of conservation, improving our understanding of species responses to ecological constraints and environmental changes over evolutionary timescales.

genomics↗

bioRxiv2024

Host-use Drives Convergent Evolution in Clownfish and Disentangles the Mystery of an Iconic Adaptive Radiation

Clownfishes (Amphiprioninae) are a fascinating example of marine radiation. From a central Pacific ancestor, they quickly colonized the coral reefs of the Indo-Pacific and diversified independently on each side of the Indo-Australian Archipelago. Their association with venomous sea anemones is often thought to be the key innovation that enabled the clownfish radiation. However, this intuition has little empirical or theoretical support given our current knowledge of the clade. To date, no ecological variable has been identified that can explain clownfish niche partitioning, phenotypic evolution, species co-occurrence, and thus, the adaptive radiation of the group. Our synthetic work solves this long-standing mystery by testing the influence of sea anemone host use on phenotypic divergence. We provide the first major revision to the known clownfish-sea anemone host associations in over 30 years, accounting for host associations in a biologically relevant way. We gathered whole-genome data for all 28 clownfish species and reconstructed a fully supported species tree for the Amphiprioninae. Integrating this new data into comparative phylogenomic approaches, we demonstrate for the first time, that the host sea anemones are the drivers of convergent evolution in clownfish color pattern and morphology. During the adaptive radiation of this group, clownfishes in different regions that associate with the same hosts have evolved the same phenotypes. Comparative genomics also reveals several genes under convergent positive selection linked to host specialisation events. Our results identify the sea anemone host as the key ecological variable that disentangles the entire adaptive radiation. As one of the most recognizable animals on the planet and an emerging model organism in the biological sciences, our findings bear on the interpretation of dozens of prior studies on clownfishes and will radically reshape research agendas for these iconic organisms.

evolutionary biology↗

bioRxiv2024

Eluding anemone nematocysts: are clownfish deprived of N-acetylated sugars in their skin mucus?

The clownfish - sea anemone system is a great example of symbiotic mutualism where host << toxicity >> does not impact its symbiont partner, although the underlying protection mechanism remains unclear. The regulation of nematocyst discharge in cnidarians involves N-acetylated sugars like sialic acid, that bind chemoreceptors on the tentacles of sea anemones, leading to the release of stings. It has been suggested that clownfish could be deprived of sialic acid on their skin surface, sparing them from being stung and facilitating mutualism with sea anemones. In this study, we sampled the skin mucus of two anemone symbionts, the clownfish Amphiprion akindynos and the juvenile damselfish Dascyllus trimaculatus, as well as two non-symbiotic adult damselfish Pomacentrus moluccensis and P. pavo. The free and total sialic acid content, including its conjugated form, and three other intermediates of this pathway were quantified using a stable isotope dilution mass spectrometry approach. We found significantly higher levels of sialic acid and its precursor in the non-symbiotic damselfishes. Concentrations of total sialic acid in anemone symbionts ranged between 13 {micro}M and 16 {micro}M, whereas the non-symbiotic damselfishes ranged between 21 {micro}M and 30 {micro}M. The presence of this metabolite and its precursors, as triggers of nematocyst discharge, in anemone symbionts, suggests that this is not the direct mechanism of protection or that the trigger is concentration dependent. This experiment demonstrates that anemone symbionts are not spared by nematocysts because of a lack of N-acetylated sugars, as previously thought.

molecular biology↗

bioRxiv2024

Ongoing hybridisation among clownfishes: the genomic architecture of the Kimbe Bay hybrid zone

Hybrid zones - locations where genetically distinct lineages interact and reproduce - are remarkable resources for exploring the evolutionary trajectory of species. Not only can we learn from hybrid zones about the mechanisms of speciation and how reproductive isolation is maintained, but we can also study their impact on evolutionary processes. Thanks to the advent of next-generation sequencing, we are now able to gain new insight into the structure of hybrid genomes and the factors influencing the outcome of hybridisation. Here, we focus on the Kimbe Bay hybrid zone, a narrow region in the Pacific Ocean where two species of clownfish - Amphiprion chrysopterus and A. sandaracinos - hybridise and give rise to the hybrid A. leucokranos. Based on whole-genome sequencing, we found that the hybrid zone is mainly composed of first-generation hybrids, the first evidence of F2 hybrids in the wild and early backcrosses with A. sandaracinos. The recurrent backcrossing with one of the parental species might lead to adaptive introgression, with few adaptive introgressed loci from A. chrysopterus integrated into the A. sandaracinos genomic background. This study builds upon the growing literature body relative to the evolutionary outcomes of hybridisation and its importance in the evolution of many species.

evolutionary biology↗

bioRxiv2023

Recurrent gene flow events shaped the diversification of the clownfish skunk complex

Clownfish (subfamily Amphiprioninae) are an iconic group of coral reef fish that evolved a mutualistic interaction with sea anemones, which was shown to have triggered the adaptive radiation of the group. Within clownfishes, the skunk complex is particularly interesting as, besides ecological speciation, gene flow between species and hybrid speciation are suggested to have shaped the diversification of the group. We investigated, for the first time, the mechanisms underlying the diversification of this complex. By taking advantage of their disjunct geographical distribution, we obtained whole-genome data of sympatric and allopatric populations of the three main species of the complex (Amphiprion akallopisos, A. perideraion and A. sandaracinos). We examined the population structure, genomic divergence patterns and introgression signals, and performed demographic modeling to identify the most realistic diversification scenario. We excluded scenarios of strict isolation, of hybrid origin of A. sandaracinos, and ruled out the presence of extensive gene flow in sympatry. We discovered moderate gene flow from A. perideraion to the ancestor of A. akallopisos + A. sandaracinos and weak gene flow between the species in the Indo-Australian Archipelago throughout the diversification process of the group. We identified introgressed regions in A. sandaracinos and detected two large regions of high divergence in A. perideraion, likely maintained by the disruption of recombination. Altogether, our results show that ancestral hybridization events shaped the groups diversification. However, more recent gene flow is less pervasive than initially thought and suggests a role of host repartition or behavioral barriers in maintaining the genetic identity of the species in sympatry.

evolutionary biology↗

bioRxiv2023

Gene flow throughout the evolutionary history of a polymorphic and generalist clownfish

Even seemingly homogeneous on the surface, the oceans display high environmental heterogeneity across space and time. Indeed, different soft barriers structure the marine environment, which offers an appealing opportunity to study various evolutionary processes such as population differentiation and speciation. Here, we focus on Amphiprion clarkii (Actinopterygii; Perciformes), the most widespread of clownfishes that exhibits the highest colour polymorphism. Clownfishes can only disperse during a short pelagic larval phase before their sessile adult lifestyle, which might limit connectivity among populations, thus facilitating speciation events. Consequently, the taxonomic status of A. clarkii has been under debate. We used whole-genome resequencing data of 67 A. clarkii specimens spread across the Indian and Pacific Oceans to characterise the species population structure, demographic history, and colour polymorphism. We found that A. clarkii spread from the Indo-Pacific Ocean to the Pacific and Indian Oceans following a stepping-stone dispersal and that gene flow was pervasive throughout its demographic history. Moreover, edge populations exhibited more similar colouration patterns compared to central populations. However, we demonstrate that colour polymorphism is not associated with population structure, thus, colour phenotype is unreliable in assessing the taxonomic status of A. clarkii. Our study further highlights the power of whole-genome comparative studies to determine the taxonomy of geographically wide-ranging and phenotypically diverse species, supporting the status of A. clarkii as a single species.

evolutionary biology↗

bioRxiv2023

Detecting diversifying selection for a trait from within and between-species genotypes and phenotypes

To quantify selection acting on a trait, methods have been developed using either within or between-species variation. However, methods using within-species variation do not integrate the changes at the macro-evolutionary scale. Conversely, current methods using between-species variation usually discard within-species variation, thus not accounting for processes at the micro-evolutionary scale. The main goal of this study is to define a neutrality index for a quantitative trait, by combining within- and between-species variation. This neutrality index integrates nucleotide polymorphism and divergence for normalizing trait variation. As such, it does not require estimation of population size nor of time of speciation for normalization. Our index can be used to seek deviation from the null model of neutral evolution, and test for diversifying selection. Applied to brain mass and body mass at the mammalian scale, we show that brain mass is under diversifying selection. Finally, we show that our test is not sensitive to the assumption that population sizes, mutation rates and generation time are constant across the phylogeny, and automatically adjust for it.

evolutionary biology↗

bioRxiv2023

Improved estimation of molecular evolution coupling stochastic simulations and deep learning

Models have always been central to inferring molecular evolution and to reconstructing phylogenetic trees. Their use typically involves the development of a mechanistic framework reflecting our understanding of the underlying biological processes, such as nucleotide substitutions, and the estimation of model parameters by maximum likelihood or Bayesian inference. However, deriving and optimizing the likelihood of the data is not always possible under complex evolutionary scenarios or tractable for large datasets, often leading to unrealistic simplifying assumptions in the fitted models. To overcome this issue, we couple stochastic simulations of genome evolution with a new supervised deep learning model to infer key parameters of molecular evolution. Our model is designed to directly analyze multiple sequence alignments and estimate per-site evolutionary rates and divergence, without requiring a known phylogenetic tree. The accuracy of our predictions matches that of likelihood-based phylogenetic inference, when rate heterogeneity follows a simple gamma distribution, but it strongly exceeds it under more complex patterns of rate variation, such as codon models. Our approach is highly scalable and can be efficiently applied to genomic data, as we show on a dataset of 26 million nucleotides from the clownfish clade. Our simulations also show that the per-site rates obtained by deep learning increase the likelihood of the true tree and could therefore lead to more accurate phylogenetic inference. We propose that future advancements in phylogenetic analysis will benefit from a semi-supervised learning approach that combines deep-learning estimation of substitution rates, which allows for more flexible models of rate variation, and probabilistic inference of the phylogenetic tree, which guarantees interpretability and a rigorous assessments of statistical support.

evolutionary biology↗