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Pie, M. R.

Publications and source records attributed to Pie, M. R..

At least 19 recordsLinked to original sources

Climate isolation and percolation as drivers of terrestrial vertebrate richness

Climate is a strong predictor of global species richness, but the effects of climatic conditions are difficult to separate from the geography of the climates themselves. Recent work in climate space has shown that the area and isolation of discrete climatic conditions explain broad-scale richness gradients, yet the internal spatial cohesion of those climates remains poorly characterized. Here, we introduce climate percolation as a complementary descriptor of climate geography, measuring the degree to which the total area of a climate bin is concentrated within effectively connected fragments. Using global range maps for amphibians, birds, mammals and reptiles, we quantified species richness across a two-dimensional climate space defined from 12 climatic variables and evaluated the independent and joint effects of climate area, climate isolation and climate percolation across multiple climate-space resolutions. Climate isolation and percolation were strongly coupled: their first joint axis explained, on average, more than 95% of their shared variation, revealing a dominant gradient of climate fragmentation along which geographically isolated climates are also internally subdivided. Despite this collinearity, percolation consistently outperformed isolation in cross-validation across all four vertebrate groups, with particularly strong predictive gains for birds and mammals. The largest improvements, however, came from the shared isolation-percolation axis, indicating that vertebrate richness in climate space is more strongly associated with the integrated geographical structure of climates than with either inter-fragment distance or internal cohesion alone. These results suggest that climate fragmentation is a multidimensional property of environmental space, combining both the distance among climate fragments and the dominance structure of connected areas. By extending climate-space approaches from area and isolation to percolation, our framework provides a more complete description of how the geography of climate may shape global richness gradients and offers a structural basis for anticipating how future changes in climate connectivity could alter biodiversity patterns.

ecology↗

Unlocking karst biodiversity with eDNA: a validated qPCR toolkit for discovery and monitoring the worlds largest cavefish radiation (Cyprinidae: Sinocyclocheilus)

Environmental DNA (eDNA) enables non-invasive detection of aquatic species through trace genetic material, but uncertainties regarding DNA persistence, detection accuracy, and the limited taxonomic resolution of stygian fauna have constrained its use in subterranean aquatic environments. The species-rich cavefish genus Sinocyclocheilus, endemic to the karst landscapes of southwestern China, offers an excellent model for understanding these issues. With over eighty species, many of which are endemics, and many still being described, this lineage is taxonomically rich, poorly understood, and highly threatened. However, access to these deep habitats remains challenging, making eDNA detection one of the most practical methods for documenting and monitoring them. We developed and validated a qPCR assay targeting a hypervariable region of the mitochondrial 16S rRNA gene, allowing for genus-specific detection and species-level identification. Laboratory and field evaluations involving 47 representative species confirmed high specificity and a detection limit of around 20 DNA copies per reaction (R{superscript 2} = 0.995). Controlled degradation studies indicated that temperature and pH strongly affect DNA persistence. Field surveys across 47 karst sites (37 caves and 10 surface sites) detected Sinocyclocheilus eDNA in only 33 caves. Dideoxy sequencing of qPCR amplicons provided species-level identification and phylogenetic validation, including detection of a new species. At three cave sites yielding multiple qPCR detections, 16S metabarcoding enabled species discrimination. By integrating qPCR assay validation, eDNA degradation modelling, hierarchical genetic identification via dideoxy sequencing, and selective NGS metabarcoding, this study presents a strategic framework for applying eDNA methods in subterranean environments.

evolutionary biology↗

Integrative Species Delimitation in a Speciation Continuum: Phylogenomics, Cryptic Diversity, Diversification and Historical Biogeography of Sinocyclocheilus Cavefish

The transition from structured populations to distinct species often unfolds along a speciation continuum. However, empirically dissecting this continuum poses challenges for species delimitation, particularly in rapid radiations marked by recent divergence, incomplete lineage sorting, and gene flow. The species-rich Sinocyclocheilus cavefish radiation of Southwest China, an emerging evolutionary multi-species model system, shows a notable mismatch between its high morphological diversity and the limited divergence observed in commonly used mitochondrial DNA markers. This pattern suggests that true species richness may be underestimated. Yet, comprehensive genome-wide approaches to resolve species boundaries in this group are still lacking. To address this, we combine phylogenomics, coalescent-based species delimitation, population genetics, and historical biogeography, using both genome-wide RAD-seq and multi-locus Sanger data (including nuclear and mitochondrial DNA). Our phylogenomic analyses resolve major clades and reveal substantial cryptic diversity, well beyond what was detected by earlier markers. Species delimitation based on RAD-seq genomic data identifies several cryptic evolutionary lineages. We recover a dynamic divergence history shaped by isolation and episodic connectivity. Biogeographic reconstruction supports a mid-Miocene origin following a major vicariance event, with subsequent founder-event dispersals into subterranean habitats. This long history of fragmentation is further complicated by reticulation and ancestral polymorphism, and is reflected in present-day patterns of restricted gene flow across river valleys. These results highlight the utility of integrative genomic approaches in resolving species boundaries and uncovering the evolutionary processes that underlie high diversity in large and complex radiations.

evolutionary biology↗

Demographic history explains mitochondrial genome capture and mitonuclear discordance among South American collared Tropidurus lizards

Mitonuclear discordance--evolutionary discrepancies between mitochondrial and nuclear DNA phylogenies--can arise from various factors, including introgression, incomplete lineage sorting, recent or ancient demographic fluctuations, sex-biased dispersal asymmetries, among others. Understanding this phenomenon is crucial for accurately reconstructing evolutionary histories, as failing to account for discordance can lead to misinterpretations of species boundaries, phylogenetic relationships, and historical biogeographic patterns. We investigate the evolutionary drivers of mitonuclear discordance in the Tropidurus spinulosus species group, which contains nine species of lizards inhabiting open tropical and subtropical environments in South America. Using a combination of population genetic and phylogenomic approaches applied to mitochondrial and nuclear data, we identified different instances of gene flow that occurred in ancestral lineages of extant species. Our results point to a complex evolutionary history marked by prolonged isolation between species, demographic fluctuations, and potential episodes of secondary contact with genetic admixture. These conditions likely facilitated mitochondrial genome capture while diluting signals of nuclear introgression. Furthermore, we found no strong evidence supporting incomplete lineage sorting or natural selection as primary drivers of the observed mitonuclear discordance. Therefore, the unveiled patterns are most consistent with neutral demographic processes, coupled with ancient mitochondrial introgression, as the main factors underlying the mismatch between nuclear and mitochondrial phylogenies in this system. Future research could further explore the role of other demographic processes, such as asymmetric sex-biased dispersal, in shaping these complex evolutionary patterns.

genomics↗

Ant head shape evolved to compromise bite-induced mechanical demands

Heads of ant workers harbour the mouthparts and respective musculature, varying drastically in morphology. The mandible adductor muscles occupy most of the heads internal volume, and their contraction generates forces that could risk cuticle failure. Here we quantified ant worker plane head shape disparity and explored how it influences stress dissipation under biting conditions. We combined a geometric morphometric approach under a phylogenetic comparative framework and biomechanical simulations to test the hypothesis that idealized head shapes poorly explored by current ant lineages exhibit biomechanical limitations that could be negatively selected, while an idealized shape representing the median ant worker head shape would present a superior mechanical performance. Our results revealed that narrow heads with deep vertex depressions distribute stresses more evenly, but usually at high levels. Broader heads with convex posterior margins, in contrast, concentrate stresses around the mandibular articulations, making them more prone to failure under high bite forces. Most ant lineages exhibit a head shape that allows mild stresses to spread along the head while concentrating higher stresses around the mandibular articulations. This evolutionary pattern results in a limited morphospace occupation, with most ants clustering around a typical shape and only a few lineages exploring extreme morphologies.

zoology↗

Caves as species pumps: key innovations, isolation, and periodic introgression drive the world's largest cavefish radiation in a dynamic karstic landscape

Species diversification is shaped by intricate interactions among biotic drivers, including gene flow, hybridization, and key innovations, and abiotic drivers, such as historical climate change, geological events, and ecological opportunity. However, the relative contributions of these drivers in large radiations remain poorly understood. We investigate the interplay among these factors in the diversification of Sinocyclocheilus, a cavefish radiation comprising 79 species. Sinocyclocheilus include typical surface-dwelling forms, with well-developed eyes and pigmentation, to cave-dwelling forms with regressed eyes, reduced pigmentation, and the presence of a horn and a hump. Using reduced representation genomic data (RADseq), we show extensive gene flow events across different species, with introgression playing a key role compared to incomplete lineage sorting in creating phylogenetic discordance and contributing genetic variation for cave adaptation and diversification in this group. Key traits such as eye degeneration, reduced pigmentation, and horn evolved independently multiple times, as adaptations for effectively exploiting cave environments. Furthermore, the uplift of the Tibetan plateau and the late Miocene cooling also significantly impacted speciation rates. Demographic analyses suggest population expansions during the Gonghe Movement and stability during the Last Glacial Maximum, possibly due to cave refugia. Periodic events of introgression promoted by isolation and reconnections due to the changing climate and geological activity, combined with the repeated evolution of key cave-adapted traits, are the primary drivers of this radiation. Our findings underscore the complex interplay of biotic and abiotic factors in the evolution of Sinocyclocheilus fish, offering new insights into the mechanisms of cave adaptation and diversification.

evolutionary biology↗

Population of the Parana Antwren Formicivora acutirostris reassessed as being 64% smaller: Revision of its conservation status and assessment of its Green Status, with management proposals

Assessing the conservation status of a species is important for designing effective conservation measures. Consequently, it is often vital to review it to update biodiversity management. Formicivora acutirostris is a bird from coastal marshes and related habitats of Brazils southern flood plains. It is considered endangered in Brazil but near threatened globally. In 2007, its area of occupancy (AOO) and population size were estimated based on assessment of aerial photographs from 1978 and 1980. Since the species is threatened and occupies a small area scattered across a coastal pressured region, we aimed to reassess its conservation status and assess its Green Status. We compiled new records, conducted new density estimates, and compared the previous mapping with orbital imagery to estimate the current AOO, habitat loss, population size, and review its conservation status based on the International Union for Conservation of Nature criteria. The species is distributed across 10 populations, including two new populations from the southern coast of Santa Catarina to the northern coast of Rio Grande do Sul. We estimated the AOO at 4,102 ha and the population as 6,284 mature territorial individuals. The previously mapped AOO has decreased by 1,535 ha due to ecological succession. The loss of habitat due to invasion by exotic grasses is the main anthropogenic impact. We recommend that the species be considered vulnerable. The Green Status indicates that the Conservation Legacy of actions taken thus far and the Conservation Dependence of ongoing actions are null, but it highlights the importance of future actions for species conservation. We propose the establishment of exotic-free zones as small geographic areas with a significant amount of minimally invaded environments, which we suggest as priority areas for the conservation of the species due to their cost-effective management potential. We also propose assisted colonization to enhance its long-term conservation.

ecology↗

Wall-following: phylogenetic context of an enhanced behaviour in stygomorphic Sinocyclocheilus (Cypriniformes: Cyprinidae) cavefishes

With 75 known species, the freshwater-fish genus Sinocyclocheilus is the largest cavefish radiation in the world, emerging as a model system for evolutionary studies. They show multiple adaptations for cave dwelling (stygomorphic adaptations), which include a range of traits such as eye degeneration (Normal-eyed, Micro-eyed and Eyeless), depigmentation of skin, and in some species, the presence of "horns". Their behavioural adaptations to subterranean environments, however, are poorly understood. Wall-following (WF) behaviour, where an organism remains in close contact with the boundary demarcating its habitat when in the dark, is a peculiar behaviour observed in a wide range of animals and is enhanced in some cave dwellers. Hence, we hypothesize wall-following to be present also in Sinocyclocheilus, possibly enhanced in Eyeless species compared to species with visual cues (Normal / Micro-eyed species). Using 13 species representative of Sinocyclocheilus radiation and eye-morphs, we designed a series of assays, based on pre-existing methods for Astyanax mexicanus behavioural experiments, to examine wall-following behaviour under three stimulation conditions. Our results indicate that eyeless species exhibit significantly enhanced levels of WF compared to Normal-eyed species, with Micro-eyed forms demonstrating intermediate levels. Using a mtDNA based dated phylogeny (chronogram with four clades A - D), we traced the degree of WF of these forms to outline common patterns. We show that intensity of WF behaviour is high in the subterranean clades (B & C) compared to clades with free-living species (A & D). Experiments on WF behaviour revealed that eyeless species are highly sensitive to vibrations, whereas normal-eyed species are the least sensitive. Since WF behaviour is present to some degree in all Sinocyclocheilus species, and given that these fishes evolved in the late Miocene, we identify this behaviour as being ancestral with WF enhancement related to cave occupation. Our results from this diversification-scale study of cavefish behaviour suggest that enhanced wall-following behaviour may be a convergent trait across all stygomorphic cavefish lineages. Significance statementSinocyclocheilus, a genus of 75 species of freshwater cavefish, is an emerging model system in evolutionary studies. Their adaptations for subterranean life, including eye degeneration, skin depigmentation, and horn-like structures, are well-known, but their behavioural adaptations remain understudied. Here we focus on a phenomenon, called "wall-following," where fish stay close to the cave walls in absence of light. We hypothesized that this behaviour would be more pronounced in eyeless species. We selected 13 species, representative of the diversity of the genus and eye types, and observed their wall-following behaviour under different conditions. Results were intriguing; eyeless species exhibited heightened wall-following behaviour compared to their sighted counterparts, with small-eyed species falling in between. Researchers also mapped this behaviour on a phylogenetic tree, discovering a pattern: cave-dwelling clades showed stronger wall-following than free-living ones. Wall-following is prevalent in all Sinocyclocheilus species and, given the evolutionary history of the genus, is considered an ancestral behaviour that intensified with cave adaptation. These findings contribute to our understanding of convergent evolution, suggesting that enhanced wall-following may be a shared trait among diverse cavefish lineages.

animal behavior and cognition↗

A frog adaptive radiation: Ecomorphological evolution in Old-World shrub frogs (Rhacophoridae: Pseudophilautus)

Ecomorphs result from divergent natural selection, leading to species-rich adaptive radiations. Identifying ecomorphs and the resulting adaptive radiations in frogs is challenging due to conserved morphology and high species diversity. In this study, we demonstrate the ecological and climate specializations that have driven the diversification of shrub frogs of the genus Pseudophilautus in Sri Lanka, a tropical continental island. We use a time-calibrated phylogeny, morphometric analyses, and climate-niche evolution, and identify five ecomorphological categories, including Tree-shrub, Rock-boulder, Leaf-litter, Habitat Generalists, and Canopy forms, and describe their evolution. Body size is the primary factor separating species, and specific body features correlate with habitat type. Ecomorphs likely evolved multiple times in disparate lineages, and in different regions and altitudes, during cold climatic periods owing to monsoon cycles resulting from the Himalayan-Tibetan orogeny. The common ancestor was a medium-sized, wet-adapted, tree-shrub habitat specialist which originated in the late Oligocene. Extreme size classes (diminutive leaf litter forms and large canopy forms) evolved recently and suggest that morphological disparity arose late in diversification, possibly aided by favorable climates. This work will facilitate understanding of adaptive radiations in frogs, which possibly will help uncover the prevalence of subtle adaptive radiations in frogs, just as in tailed-vertebrates.

evolutionary biology↗

Substantial variation in species ages among vertebrate clades

Ecological and evolutionary studies traditionally assume that species are comparable units of biodiversity. However, not only this assumption is rarely tested, but also there have been few attempts even to assess variation in most emergent, species-level traits and their corresponding underlying mechanisms. One such trait is species age, here defined as the time since the most recent common ancestor between a given species and its sister lineage. In this study, we demonstrate that different terrestrial vertebrate clades vary considerably in the age of their constituent species. In particular, species ages were youngest in mammals and birds as opposed to squamates and amphibians, although considerable variation was found within those clades as well. Sensitivity analyses showed that these results are unaffected by phylogenetic uncertainty or incomplete taxonomic sampling. Interestingly, there was little geographical correspondence in mean species age across taxa, as well as with temperature and precipitation stability over the past 21,000 years. We discuss candidate mechanisms that might explain differences in species ages among clades, and explore the implications of these findings in relation to recent advances in age-dependent speciation and extinction models of diversification.

evolutionary biology↗

Diurnality shapes the visual opsin genes of colorful Neotropical frogs

Amphibians are ideal for studying visual system evolution because their biphasic (aquatic and terrestrial) life history and ecological diversity expose them to a broad range of visual conditions. Here we evaluate signatures of selection on visual opsin genes across Neotropical anurans and focus on three diurnal clades that are well-known for the concurrence of conspicuous colors and chemical defense (i.e., aposematism): poison frogs (Dendrobatidae), Harlequin toads (Bufonidae: Atelopus), and pumpkin toadlets (Brachycephalidae: Brachycephalus). We found evidence of positive selection on 44 amino acid sites in LWS, SWS1, SWS2, and RH1 opsin genes, of which one in LWS and two in RH1 have been previously identified as spectral tuning sites in other vertebrates. Given that anurans have mostly nocturnal habits, the patterns of selection revealed new sites that might be important in spectral tuning for frogs, potentially for adaptation to diurnal habits and for color-based intraspecific communication. Furthermore, we provide evidence that SWS2, normally expressed in rod cells in amphibians, has likely been lost in the ancestor of Dendrobatidae, suggesting that under low-light levels, dendrobatids have inferior wavelength discrimination compared to other frogs. This loss might follow the origin of diurnal activity in dendrobatids and could have implications for their chemical ecology, biodiversity, and behavior. Our analyses show that assessments of opsin diversification in understudied groups could expand our understanding of the role of sensory system evolution in ecological adaptation.

evolutionary biology↗

Climatic niche evolution of infectious diseases driving amphibian declines

Climate change and infectious diseases continue to drive global amphibian population declines, contributing to one of the greatest vertebrate extinctions of the Anthropocene. Currently around 16% amphibian species across the world are affected by four pathogens - Batrachochytrium dendrobatidis (Bd), B. salamandrivorans (Bsal), Ranavirus and Perkinsea. A climatic context behind the dispersal of some of these diseases is hypothesized. However, the interplay between niche conservatism (NC) and climatic niche evolution (CNE), essential to understand disease evolution and dispersal, has so far received little attention. Here we show that the impacts of amphibian pathogens are intensifying as their climatic niches evolve. NC-based analyses suggest that niches of these diseases overlap, especially in Europe and East/southeast Asia (ESEA), and that all four pathogens will continue to devastate amphibians through seasonality shifts and range expansions, penetrating deeper into temperate regions and global amphibian diversity hotspots. Bd will spread over diversity-rich mountain ranges and ranaviruses will overwhelm lowlands. CNE-based analyses suggest that the earliest lineages of these diseases originated in colder regions and that some lineages subsequently evolved towards warmer climatic niches. We caution that quiescent, warm-adapted strains are likely to become widespread and novel ranaviruses adapted to local climatic conditions and new hosts are likely to emerge. These results portend the dangers of introducing pathogens into new regions given their ability to adapt to changing climate scenarios. In a climatic background conducive to most of these diseases, frequent monitoring, enhanced biosecurity measures and policy reforms are needed for disease control and mitigation.

evolutionary biology↗

Wanted not, wasted not: Searching for non-target taxa in environmental DNA metabarcoding by-catch

Metabarcoding of environmental DNA is based on primers that are specific to the target taxa (e.g. bacteria, zooplankton, fishes). However, due to the nature of the commonly used protocols, regardless of the chosen primers, several sequences of non-target species will inevitably be generated, but are usually discarded in commonly used bioinformatics pipelines. These non-target sequences might contain important biological information about the presence of other species in the studied habitats and its potential for ecological studies is still poorly understood. Here, we analyzed the presence of mammal and bird species in aquatic environmental samples that were originally amplified targeting teleost fish species. After all cleaning and checking steps, we kept 21 amplicon sequence variants (ASVs) belonging to mammals and ten to birds. Most ASVs were taxonomic assigned to farm/domestic animals, such as cats, cows, and ducks. Yet, we were able to identify a native semi-aquatic mammal, the capybara, in the samples. Four native bird species and a non-native potentially invasive bird (Corvus sp.) were also detected. Although the data derived from these samples for mammals and birds are of limited use for diversity analyses, our results demonstrate the potential of aquatic samples to characterize non-aquatic birds and highlight the presence of a potentially invasive species that had not been recorded before in the region.

ecology↗

Evolving in the darkness: phylogenomics of Sinocyclocheilus cavefishes highlights recent diversification and cryptic diversity

Troglomorphism-- any morphological adaptation enabling life to the constant darkness of caves, such as loss of pigment, reduced eyesight or blindness, over-developed tactile and olfactory organs--has long intrigued biologists. However, inferring the proximate and ultimate mechanisms driving the evolution of troglomorphism in freshwater fish requires a sound understanding of the evolutionary relationships between surface and troglomorphic lineages. We use Restriction Site Associated DNA Sequencing (RADseq) to better understand the evolution of the Sinocyclocheilus fishes of China. With a remarkable array of derived troglomorphic traits, they comprise the largest cavefish diversification in the world, emerging as a multi-species model system to study evolutionary novelty. We sequenced a total of 120 individuals throughout the Sinocyclocheilus distribution. The data comprised a total of 646,497bp per individual, including 4378 loci and 67,983 SNPs shared across a minimum of 114 individuals at a given locus. Phylogenetic analyses using either the concatenated RAD loci (RAxML) or the SNPs under a coalescent model (SVDquartets, SNAPP) showed a high degree of congruence with similar topologies and high node support (> 95 for most nodes in the phylogeny). The major clades recovered conform to a pattern previously established using Sanger-based mt-DNA sequences, with a few notable exceptions. We now recognize six major clades in this group, elevating the blind cavefish S. tianlinensis and the micro-eyed S. microphthalmus as two new distinct clades due to their deep divergence from other clades. PCA plots of the SNP data also supports the recognition of six major clusters of species congruent with the identified clades based on the spatial arrangement and overlap of the species in the PC space. A Bayes factor delimitation (BFD) analysis showed support for 21 species, recognizing 19 previously described species and two putative new cryptic ones. Two species whose identities were previously disputed, S. furcodorsalis and S. tianeensis, are supported here as distinct species. In addition, our multi-species calibrated tree in SNAPP suggests that the genus Sinocyclocheilus originated around 10.5 Mya, with most speciation events occurring in the last 2 Mya, likely favored by the uplift of the Qinghai-Tibetan Plateau and cave occupation induced by climate-driven aridification during this period. These results provide a firm basis for future comparative studies on the evolution of Sinocyclocheilus and its adaptations to cave life.

evolutionary biology↗

Drivers of an epic radiation: the role of climate and islands in species diversification and reproductive-mode evolution of Old-World tree frogs

Although large diversifications of species occur unevenly across space and evolutionary lineages, the relative importance of their driving mechanisms, such as climate, ecological opportunity and key innovations, remains poorly understood. Here, we explore the remarkable diversification of rhacophorid frogs, which represent six percent of global amphibian diversity, utilize four distinct reproductive modes, and span a climatically variable area across mainland Asia, associated continental islands, and Africa. Using a complete species-level phylogeny, we find near-constant diversification rates but a highly uneven distribution of species richness. Montane regions on islands and some mainland regions have higher phylogenetic diversity and unique assemblages of taxa; we identify these as cool-wet refugia. Starting from a centre of origin, rhacophorids reached these distant refugia by adapting to new climatic conditions ( niche evolution-dominant), especially following the origin of key innovations such as terrestrial reproduction (in the Late Eocene) or by dispersal during periods of favourable climate ( niche conservatism-dominant).

evolutionary biology↗

Monitoring fish communities through DNA metabarcoding in the fish pass system of the second largest hydropower plant in the world

The Itaipu Hydroelectric Power Plant is the second largest in the world in power generation. The artificial barrier created by its dam imposes an obstacle for fish migration. Thus, in 2002, a fish pass system, named Piracema Channel, was built to allow fish to access areas upstream of the reservoir. We tested the potential of environmental DNA metabarcoding to monitor the impact of both the dam and associated fish pass system in the Parana River fish communities and to compare it with traditional monitoring methods. Using a fragment of the 12S gene, we characterized richness and community composition based on amplicon sequence variants, operational taxonomic units, and zero-radius OTUs. We combined GenBank and in-house data for taxonomic assignment. We found that different bioinformatics approaches showed similar results. Also, we found a decrease in fish diversity from 2019 to 2020 probably due to the recent extreme drought experienced in southeastern Brazil. The highest alpha diversity was recorded in the mouth of the fish pass system, located in a protected valley with the highest environmental heterogeneity. Despite the clear indication that the reference databases need to be continuously improved, our results demonstrate the analytical efficiency of the metabarcoding to monitor fish species.

ecology↗

Adapting insect caste concepts to the demands of bio-ontologies.

AbstractTerminological and conceptual ambiguity surrounding core entities such as colonies and castes continues to hinder comparative analysis, data integration, and ontological interoperability in social insect research. While extensive work has addressed the descriptive and functional aspects of social organization, less attention has been paid to the ontological status of these entities and the consequences this has for formal representation in bio-ontologies. In this study, I provide a realist ontological analysis of colonies and castes grounded in distinctions between natural units, natural kinds, and realizable entities. I argue that insect colonies constitute bona fide biological individuals (natural units) demarcated through relations of causal unity and historical continuity, and therefore suitable for treatment as unified causal systems within ontology frameworks. By contrast, I show that castes fail to satisfy the criteria for either natural units or natural kinds, due to their lack of independent causal cohesion, developmental plasticity, temporal variability, and context sensitivity. Treating castes as autonomous entities or essentialized kinds introduces systematic instability and misrepresentation in bio-ontological models. To address these issues, I propose modeling castes as realizable entities, specifically as roles instantiated in individual organ-isms and realized in biological processes under colony-regulated developmental and ecological conditions. This approach accommodates morphological canalization, phenotypic plasticity, temporal polyethism, and cross-taxon variability within a unified framework, while remaining consistent with the Basic Formal Ontology (BFO). My analysis clarifies longstanding conceptual ambiguities in the representation of social insect organization and provides a flexible, biologically faithful foundation for ontology design, supporting robust data interoperability and future empirical refinement.

zoology↗

The diversification of termites: inferences from a complete species-level phylogeny

Termites play a major role in a variety of ecological processes in tropical and subtropical biomes worldwide, such as decomposition, soil formation and aeration, and nutrient cycling. These important ecosystem services were achieved through their highly complex societies and remarkable adaptations, including the evolution of sterile worker castes, the acquisition of endosymbionts, and the capacity for extensive environmental engineering, yet the causes and consequences of their ecological success are still poorly understood. The goals of our study were (1) to provide the first complete, species-level phylogeny of all currently recognized termite species by integrating the available genetic and taxonomic data, as well as methods of phylogenetic imputation and divergence time estimation; and (2) to explore variation in speciation rates among termite lineages. We provide the inferred relationships as a set of 1,000 pseudo-posterior trees, which can be used in future comparative analyses. We demonstrate that speciation rates have been relatively constant throughout the history of termites, with two positive shifts in speciation rates: one at their origin of Euisoptera and the other concordant with evolution of Termitidae. On the other hand, there was no obvious trend towards deceleration in speciation rates for termites as a whole, nor within the most species-rich families. The provided trees might represent a valuable resource for termite comparative studies by summarizing the available phylogenetic information, while accounting for uncertainty in the inferred topologies.

evolutionary biology↗