Phylogenetic Comparative Approach Research Articles

Palms of the past: Can morphometric phytolith analysis inform deep time evolution and palaeoecology of Arecaceae?

Palm fossils are often used as evidence for warm and wet palaeoenvironments, reflecting the affinities of most modern palms. However, several extant palm lineages tolerate cool and/or arid climates, making a clear understanding of the taxonomic composition of ancient palm communities important for reliable palaeoenvironmental inference. However, taxonomically identifiable palm fossils are rare and often confined to specific facies. Although the resolution of taxonomic information they provide remains unclear, phytoliths (microscopic silica bodies) provide a possible solution because of their high preservation potential under conditions where other plant fossils are scarce. We thus evaluate the taxonomic and palaeoenvironmental utility of palm phytoliths. We quantified phytolith morphology of 97 modern palm and other monocot species. Using this dataset, we tested the ability of five common discriminant methods to identify nine major palm clades. We then compiled a dataset of species' climate preferences and tested if they were correlated with phytolith morphology using a phylogenetic comparative approach. Finally, we reconstructed palm communities and palaeoenvironmental conditions at six fossil sites. Best performing models correctly identified phytoliths to their clade of origin only 59% of the time. Although palms were generally distinguished from non-palms, few palm clades were highly distinct, and phytolith morphology was weakly correlated with species' environmental preferences. Reconstructions at all fossil sites suggested that palm communities were dominated by Trachycarpeae and Areceae, with warm, equable climates, and high, potentially seasonal rainfall. However, fossil site reconstructions had high uncertainty and often conflicted with other climate proxies. While phytolith morphology provides some distinction among palm clades, caution is warranted. Unlike prior spatially-restricted studies, our geographically and phylogenetically broad study indicates phytolith morphology may not reliably differentiate most palm taxa in deep time. Nevertheless, it reveals distinct clades, including some likely to be palaeoenvironmentally informative.

Ecological predictors of organelle genome evolution: Phylogenetic correlations with taxonomically broad, sparse, unsystematized data.

Comparative analysis of variables across phylogenetically linked observations can reveal mechanisms and insights in evolutionary biology. As the taxonomic breadth of the sample of interest increases, challenges of data sparsity, poor phylogenetic resolution, and complicated evolutionary dynamics emerge. Here, we investigate a cross-eukaryotic question where all these problems exist: which organismal ecology features are correlated with gene retention in mitochondrial and chloroplast DNA (organelle DNA or oDNA). Through a wide palette of synthetic control studies, we first characterize the specificity and sensitivity of a collection of parametric and non-parametric phylogenetic comparative approaches to identify relationships in the face of such sparse and awkward datasets. This analysis is not directly focused on oDNA, and so provides generalizable insights into comparative approaches with challenging data. We then combine and curate ecological data coupled to oDNA genome information across eukaryotes, including a new semi-automated approach for gathering data on organismal traits from less systematized open-access resources including encyclopedia articles on species and taxa. The curation process also involved resolving several issues with existing datasets, including enforcing clade-specificity of several ecological features and fixing incorrect annotations. Combining this unique dataset with our benchmarked comparative approaches, we confirm support for several known links between organismal ecology and organelle gene retention, identify several previously unidentified relationships constituting possible ecological contributors to oDNA genome evolution, and provide support for a recently hypothesized link between environmental demand and oDNA retention. We, with caution, discuss the implications of these findings for organelle evolution and of this pipeline for broad comparative analyses in other fields.

Accurate Detection of Convergent Mutations in Large Protein Alignments With ConDor.

Evolutionary convergences are observed at all levels, from phenotype to DNA and protein sequences, and changes at these different levels tend to be correlated. Notably, convergent mutations can lead to convergent changes in phenotype, such as changes in metabolism, drug resistance, and other adaptations to changing environments. We propose a two-component approach to detect mutations subject to convergent evolution in protein alignments. The "Emergence" component selects mutations that emerge more often than expected, while the "Correlation" component selects mutations that correlate with the convergent phenotype under study. With regard to Emergence, a phylogeny deduced from the alignment is provided by the user and is used to simulate the evolution of each alignment position. These simulations allow us to estimate the expected number of mutations in a neutral model, which is compared to the observed number of mutations in the data studied. In Correlation, a comparative phylogenetic approach, is used to measure whether the presence of each of the observed mutations is correlated with the convergent phenotype. Each component can be used on its own, for example Emergence when no phenotype is available. Our method is implemented in a standalone workflow and a webserver, called ConDor. We evaluate the properties of ConDor using simulated data, and we apply it to three real datasets: sedge PEPC proteins, HIV reverse transcriptase, and fish rhodopsin. The results show that the two components of ConDor complement each other, with an overall accuracy that compares favorably to other available tools, especially on large datasets.

Exploring the relationship between environment and brain morphology in anurans: a comparative phylogenetic approach

Abstract The correlation of ecological and behavioural characteristics with morphological features of brain structures has been recurrently corroborated for different vertebrate taxa, such as mammals, birds, and fishes. Here, we asked whether such correlations can also be detected in amphibians, an animal group of great interest due to its great behavioural and ecological diversity. We analysed the shape of two brain regions, cerebellum and optic tectum, of 67 species from 32 families of the order Anura (frogs and toads), applying 2D geometric morphometrics in a phylogenetic context. We found species that share similar environments and daily patterns have a similarly shaped cerebellum and optic tectum, with diurnal and arboreal species with more varied brain shapes. We did not find a correlation between activity mode or lifestyle with shape for both brain regions; however, we found that shared ancestry had an influence in their evolution. Future studies including a higher number of species from each ecological group, and from more families, would help to better clarify whether the patterns found here are a general rule for anurans.

The evolution of positive allometry and exaggerated traits in a diverse beetle clade

Abstract Understanding variation in trait allometry and the drivers of variation in sexually selected traits is a major theme in evolutionary biology. The static allometries of sexually selected traits are often positive due to disproportional investment by large individuals into trait size. Comparative studies of weapon allometry are rare, and typically focus on variation along a single dimension (e.g. length), despite weapons often diverging dramatically in shape. Furthermore, studies predominantly focus on groups where weapons have evolved as novel structures (e.g. horns), despite most weapons arising from modifications of existing structures. We test the hypothesis that sexual selection drives variation in static allometry using a phylogenetic comparative approach with 59 brentine weevil species showing remarkable variation in trait exaggeration and sexual size dimorphism (SSD). Contrasting two dimensions of head size exaggeration (a sexually selected trait) and elytron length (a nonsexual trait), positive static allometry was common across both measures of male head size, but not females, and there was positive covariance between male head static allometry and SSD across species. Positive static allometry for male elytron length was less common, but did show positive covariance with SSD, possibly due to the function of wings as compensatory structures for bearing large weapons. Our findings support the hypothesis that sexual selection drives the evolution of allometries both within and among species, and that the modification of existing structures to be used as weapons does not limit the potential for positive static allometry. Read the free Plain Language Summary for this article on the Journal blog.

Phylogeny, Ecology, and Gene Families Covariation Shaped the Olfactory Subgenome of Rodents.

Olfactory receptor (OR) genes represent the largest multigenic family in mammalian genomes and encode proteins that bind environmental odorant molecules. The OR repertoire is extremely variable among species and is subject to many gene duplications and losses, which have been linked to ecological adaptations in mammals. Although they have been studied on a broad taxonomic scale (i.e., placental), finer sampling has rarely been explored in order to better capture the mechanisms that drove the evolution of the OR repertoire. Among placental mammals, rodents are well-suited for this task, as they exhibit diverse life history traits, and genomic data are available for most major families and a diverse array of lifestyles. In this study, 53 rodent published genomes were mined for their OR subgenomes. We retrieved more than 85,000 functional and pseudogene OR sequences that were subsequently classified into phylogenetic clusters. Copy number variation among rodents is similar to that of other mammals. Using our OR counts along with comparative phylogenetic approaches, we demonstrated that ecological niches such as diet, period of activity, and a fossorial lifestyle strongly impacted the proportion of OR pseudogenes. Within the OR subgenome, phylogenetic inertia was the main factor explaining the relative variations of the 13 OR gene families. However, a striking exception was a convergent 10-fold expansion of the OR family 14 among the phylogenetically divergent subterranean mole-rat lineages belonging to Bathyergidae and Spalacidae families. This study illustrates how the diversity of the OR repertoire has evolved among rodents, both shaped by selective forces stemming from species life history traits and neutral evolution along the rodent phylogeny.

Lack of evidence for coevolution between oxytocin receptor N-terminal variants and monogamy in placental mammals

Oxytocin (OXT) is a neurohypophyseal hormone that influences a wide range of affiliative behaviors, such as pair-bonding and infant care, across mammals. The effects of OXT depend significantly on an adequate interaction with its receptor, OXTR. OXTR belongs to the G-protein coupled receptor family. The extracellular N-terminal domain of OXTR interacts with the linear C-terminal tail of OXT and is required for OXT binding. Across mammalian species there is a genetic diversity in OXTR terminal sequence. Previous work on primates has shown an association between OXTR phylogeny and monogamy. However, it is not clear whether this variation coevolved with either mating system (monogamy) or infant care behaviors (such as allomaternal care). Here, we take a phylogenetic comparative and evolutionary modeling approach across a wide range of placental mammals (n = 60) to test whether OXTR N-terminal variants co-evolved with either monogamy or allomaternal care behaviors. Our results indicate that the diversity in OXTR N-terminal region is unlikely to provide the underlying genetic bases for variation in mating system and/or allomaternal behavior as we find no evidence for co-evolution between protein sequence and affiliative behaviors. Hence, the role played by OXT in influencing affiliative behaviors is unlikely to be mediated by the genetic diversity of its receptor.

Evolution End Classification of tfd Gene Clusters Mediating Bacterial Degradation of 2,4-Dichlorophenoxyacetic Acid (2,4-D).

The tfd (tfdI and tfdII) are gene clusters originally discovered in plasmid pJP4 which are involved in the bacterial degradation of 2,4-dichlorophenoxyacetic acid (2,4-D) via the ortho-cleavage pathway of chlorinated catechols. They share this activity, with respect to substituted catechols, with clusters tcb and clc. Although great effort has been devoted over nearly forty years to exploring the structural diversity of these clusters, their evolution has been poorly resolved to date, and their classification is clearly obsolete. Employing comparative genomic and phylogenetic approaches has revealed that all tfd clusters can be classified as one of four different types. The following four-type classification and new nomenclature are proposed: tfdI, tfdII, tfdIII and tfdIV(A,B,C). Horizontal gene transfer between Burkholderiales and Sphingomonadales provides phenomenal linkage between tfdI, tfdII, tfdIII and tfdIV type clusters and their mosaic nature. It is hypothesized that the evolution of tfd gene clusters proceeded within first (tcb, clc and tfdI), second (tfdII and tfdIII) and third (tfdIV(A,B,C)) evolutionary lineages, in each of which, the genes were clustered in specific combinations. Their clustering is discussed through the prism of hot spots and driving forces of various models, theories, and hypotheses of cluster and operon formation. Two hypotheses about series of gene deletions and displacements are also proposed to explain the structural variations across members of clusters tfdII and tfdIII, respectively. Taking everything into account, these findings reconstruct the phylogeny of tfd clusters, have delineated their evolutionary trajectories, and allow the contribution of various evolutionary processes to be assessed.

Physiological cold tolerance evolves faster than climatic niches in plants.

Understanding how plants respond to thermal stress is central to predicting plant responses and community dynamics in natural ecosystems under projected scenarios of climate change. Although physiological tolerance is suggested to evolve slower than climatic niches, this comparison remains to be addressed in plants using a phylogenetic comparative approach. In this study, we compared i) the evolutionary rates of physiological tolerance to extreme temperatures with ii) the corresponding rates of climatic niche across three major vascular plant groups. We further accounted for the potential effects of hardening when examining the association between physiological and climatic niche rates. We found that physiological cold tolerance evolves faster than heat tolerance in all three groups. The coldest climatic-niche temperatures evolve faster than the warmest climatic-niche temperatures. Importantly, evolutionary rates of physiological cold tolerance were faster than rates of change in climatic niches. However, an inverse association between physiological cold tolerance and responding climatic niche for plants without hardening was detected. Our results indicated that plants may be sensitive to changes in warmer temperatures due to the slower evolutionary rates of heat tolerance. This pattern has deep implications for the framework that is being used to estimate climate-related extinctions over the upcoming century.

Ecological correlates of cranial evolution in the megaradiation of dipsadine snakes

BackgroundDipsadine snakes represent one of the most spectacular vertebrate radiations that have occurred in any continental setting, with over 800 species in South and Central America. Their species richness is paralleled by stunning ecological diversity, ranging from arboreal snail-eating and aquatic eel-eating specialists to terrestrial generalists. Despite the ecological importance of this clade, little is known about the extent to which ecological specialization shapes broader patterns of phenotypic diversity within the group. Here, we test how habitat use and diet have influenced morphological diversification in skull shape across 160 dipsadine species using micro-CT and 3-D geometric morphometrics, and we use a phylogenetic comparative approach to test the contributions of habitat use and diet composition to variation in skull shape among species.ResultsWe demonstrate that while both habitat use and diet are significant predictors of shape in many regions of the skull, habitat use significantly predicts shape in a greater number of skull regions when compared to diet. We also find that across ecological groupings, fossorial and aquatic behaviors result in the strongest deviations in morphospace for several skull regions. We use simulations to address the robustness of our results and describe statistical anomalies that can arise from the application of phylogenetic generalized least squares to complex shape data.ConclusionsBoth habitat and dietary ecology are significantly correlated with skull shape in dipsadines; the strongest relationships involved skull shape in snakes with aquatic and fossorial lifestyles. This association between skull morphology and multiple ecological axes is consistent with a classic model of adaptive radiation and suggests that ecological factors were an important component in driving morphological diversification in the dipsadine megaradiation.

Coevolutionary maintenance of forked tails and song in hirundines (Aves: Hirundininae)

Both conspicuous plumage ornamentation and song are well‐known examples of sexually selected traits, but their interrelationship is not well known, perhaps in part because of confounding factors, including interspecific variation in ecology, habitat, morphology, and type of ornamentation. Here, using a phylogenetic comparative approach and the 69 species with available information (i.e. 96% of all 72 species), I examined the evolutionary relationship between forked tails and the presence/absence of song in hirundines (Aves: Hirundininae). Hirundines have similar ecology (e.g. aerial insectivores, social monogamy, and biparental provisioning), morphology (e.g. syrinx with nearly complete bronchial rings), and plumage ornamentation (i.e. a sexually selected forked tail), which provides a unique opportunity to examine the evolutionary associations between plumage ornamentation and song. In particular, hirundines have repeatedly lost their ornamentation, forked tails, setting up a condition to test their association with the evolutionary gain/loss of their simple song. After controlling for phylogeny and covariates, I demonstrated that song was less likely to be found in species with forkless tails than in species with forked tails. Two correlates of tail shape, sexual dimorphism in the overall plumage characteristics as a well‐known proxy of sexual selection and incubation type as a measure of extrapair mating opportunity, were not positively related to the presence or absence of song, indicating the importance of forked tails, rather than their correlates. The analysis of the correlated evolution of discrete characters further supported the correlated evolution of the two traits, in which forked tails and song are maintained together and less likely to be lost under the presence of each other. The current study provided macroevolutionary support for the integrated use of visual and acoustic courtship traits.

Functional Trait Variation Along Animal Invasion Pathways

Functional trait–based mediation of animal invasions is only now developing, yet it is already showing as much promise as the approach has for plant invasion biology. Here, we provide a theory-founded examination of functional trait–based ecology with respect to animal invasions, together with a review of the empirical research. Recent developments in the scaling of traits to ecosystems, along with the frameworks for invasion ecology, provide a powerful foundation for investigations. Despite growing empirical research in this field, interspecific comparisons incorporating comparative phylogenetic approaches remain uncommon, and those examining assemblages are even more so. Despite the importance of time since introduction, it is rarely included in studies. Broad environmental tolerances, life history traits associated with fast growth or greater reproduction, and in some cases, greater phenotypic plasticity of traits, mediate the success of animal alien species at various stages of invasion, although introduction effort remains one of the most important factors affecting success.

Brain size and shape diversification in a highly diverse South American clade of rodents (Echimyidae): a geometric morphometric and comparative phylogenetic approach

Abstract Brain morphological variation is analysed through virtual endocasts in a highly diversified clade of caviomorph rodents belonging to the family Echimyidae. Diversification in brain size and shape is explored through geometric morphometrics and comparative phylogenetic analyses. The results indicate that brain shape is largely independent of general size and reveal different trends in brain size and shape. Fossorial Euryzygomatominae, arboreal Echimyini and the semi-aquatic Myocastorini Myocastor show high encephalization; the former with a greater contribution from the olfactory bulb and petrosal lobe, and the latter two with a larger surface area of neocortex. The Euryzygomatomyinae and Myocastorini of terrestrial habits show low encephalization with a low contribution of the neocortex. Phylogenetic comparative analyses suggest that endocranial morphological evolution would have been influenced by both phylogeny and locomotor habits. The concurrence of the best fit of the Ornstein–Uhlenbeck model and the significant phylogenetic signal in the datasets suggests the involvement of constraints on morphological diversification within the major clades, as expected under phylogenetic conservatism. This could be explained by an early establishment of a particular endocranial morphology in each major clade, which would have been maintained with relatively little change.

Phylogenetic comparative approach reveals evolutionary conservatism, ancestral composition, and integration of vertebrate gut microbiota.

How host-associated microbial communities evolve as their hosts diversify remains equivocal: How conserved is their composition? What was the composition of ancestral microbiota? Do microbial taxa covary in abundance over millions of years? Multivariate phylogenetic models of trait evolution are key to answering similar questions for complex host phenotypes, yet they are not directly applicable to relative abundances, which usually characterize microbiota. Here, we extend these models in this context, thereby providing a powerful approach for estimating phylosymbiosis (the extent to which closely related host species harbor similar microbiota), ancestral microbiota composition, and integration (evolutionary covariations in bacterial abundances). We apply our model to the gut microbiota of mammals and birds. We find significant phylosymbiosis that is not entirely explained by diet and geographic location, indicating that other evolutionary-conserved traits shape microbiota composition. We identify main shifts in microbiota composition during the evolution of the two groups and infer an ancestral mammalian microbiota consistent with an insectivorous diet. We also find remarkably consistent evolutionary covariations among bacterial orders in mammals and birds. Surprisingly, despite the substantial variability of present-day gut microbiota, some aspects of their composition are conserved over millions of years of host evolutionary history.

Low morphological disparity and decelerated rate of limb size evolution close to the origin of birds.

The origin of birds from theropod dinosaurs involves many changes in musculoskeletal anatomy and epidermal structures, including multiple instances of convergence and homology-related traits that contribute to the refinement of flight capability. Changes in limb sizes and proportions are important for locomotion (for example, the forelimb for bird flight); thus, understanding these patterns is central to investigating the transition from terrestrial to volant theropods. Here we analyse the patterns of morphological disparity and the evolutionary rate of appendicular limbs along avialan stem lineages using phylogenetic comparative approaches. Contrary to the traditional wisdom that an evolutionary innovation like flight would promote and accelerate evolvability, our results show a shift to low disparity and decelerated rate near the origin of avialans that is largely ascribed to the evolutionarily constrained forelimb. These results suggest that natural selection shaped patterns of limb evolution close to the origin of avialans in a way that may reflect the winged forelimb 'blueprint' associated with powered flight.

Prevalence and Molecular Characterisation of Blastocystis sp. Infecting Free-Ranging Primates in Colombia.

Infection with Blastocystis sp. has been reported in free-living and captive non-human primates (NHPs); however, surveys on Blastocystis sp. from north-western South America are scarce. This study aimed to identify Blastocystis sp. in free-ranging NHPs living in Colombia. A total of 212 faecal samples were collected from Ateles hybridus, Cebus versicolor, Alouatta seniculus, Aotus griseimembra, Sapajus apella, and Saimiri cassiquiarensis. Smears and flotation were used for morphological identification. For samples microscopically classified as positive for Blastocystis sp., we used conventional PCR to amplify and sequence two regions of the SSU rRNA gene and used Maximum Likelihood methods and Median Joining Network analyses for phylogenetic analyses. Via microscopy, 64 samples were Blastocystis sp. positive. Through molecular analyses, 18 sequences of Blastocystis sp. subtype 8 (ST8) were obtained. Strain and allele assignment together with a comparative phylogenetic approach confirmed that the sequences were ST8. Alleles 21, 156, and 157 were detected. Median Joining network analyses showed one highly frequent haplotype shared by specimens from Colombia and Peru and close relationships between haplotypes circulating in NHPs from Colombia, Ecuador, Brazil, and Mexico. This survey could support the elaboration of a more accurate epidemiological picture of the Blastocystis sp. infecting NHPs.

Ecomorphological correlates of inner and middle ear anatomy within phyllostomid bats.

Echolocation is the primary sense used by most bats to navigate their environment. However, the influence of echolocating behaviors upon the morphology of the auditory apparatus remains largely uninvestigated. While it is known that middle ear ossicle size scales positively with body mass across mammals, and that peak call frequency scales negatively with body mass among bats, there are still large gaps in our understanding of the degree to which allometry or ecology influences the morphology of the chiropteran auditory apparatus. To investigate this, we used μCT datasets to quantify three morphological components of the inner and middle ear: ossicle size, ossicle shape, and cochlear spirality. These data were collected across 27 phyllostomid species, spanning a broad range of body sizes, habitats, and dietary categories, and the relationships between these variables and ear morphology were assessed using a comparative phylogenetic approach. Ossicle size consistently scaled with strong negative allometry relative to body mass. Cochlear spirality was significantly (p=.025) associated with wing aspect ratio (a proxy for habitat use) but was not associated with body mass. From a morphological perspective, the malleus and incus exhibited some variation in kind with diet and call frequency, while stapes morphology is more closely tied to body size. Future work will assess these relationships within other chiropteran lineages, and investigate potential morphological differences in the middle and inner ear of echolocating-vs-non-echolocating taxa.

Mode and Tempo of 3D Genome Evolution in Drosophila.

Topologically associating domains (TADs) are thought to play an important role in preventing gene misexpression by spatially constraining enhancer-promoter contacts. The deleterious nature of gene misexpression implies that TADs should, therefore, be conserved among related species. Several early studies comparing chromosome conformation between species reported high levels of TAD conservation; however, more recent studies have questioned these results. Furthermore, recent work suggests that TAD reorganization is not associated with extensive changes in gene expression. Here, we investigate the evolutionary conservation of TADs among 11 species of Drosophila. We use Hi-C data to identify TADs in each species and employ a comparative phylogenetic approach to derive empirical estimates of the rate of TAD evolution. Surprisingly, we find that TADs evolve rapidly. However, we also find that the rate of evolution depends on the chromatin state of the TAD, with TADs enriched for developmentally regulated chromatin evolving significantly slower than TADs enriched for broadly expressed, active chromatin. We also find that, after controlling for differences in chromatin state, highly conserved TADs do not exhibit higher levels of gene expression constraint. These results suggest that, in general, most TADs evolve rapidly and their divergence is not associated with widespread changes in gene expression. However, higher levels of evolutionary conservation and gene expression constraints in TADs enriched for developmentally regulated chromatin suggest that these TAD subtypes may be more important for regulating gene expression, likely due to the larger number of long-distance enhancer-promoter contacts associated with developmental genes.

Genetic diversity analysis and phylogeography of Osteochilus spilurus (Cyprinidae: Labeoninae) from Bangka, Belitung, and Kalimantan Islands using Cytochrome b gene

Abstract. Kurniawan A, Hariati AM, Kurniawan A, Kurniawan N, Nugroho TW, Bidayani E, Syarif AF, wiadnya DGR. 2022. Genetic diversity analysis and phylogeography of Osteochilus spilurus (Cyprinidae: Labeoninae) from Bangka, Belitung, and Kalimantan Islands using Cytochrome b gene. Biodiversitas 23: 4738-4746. The Sundaland fish species, Osteochilus spilurus is only utilized commercially in the Belitung Islands and has yet to be domesticated. This species can characterize freshwater fish phylogeography because have a minimum of human influence. This study aimed to analyze the genetic diversity and phylogeography of O. spilurus from the Bangka, Belitung, and Kalimantan Islands using Cytochrome b gene. Twenty fish samples were collected from the Sebangau and Kapuas Rivers in Kalimantan, the Lenggang River in Belitung Island, and the Lebak River in Bangka Island. Forward LA-cyp and reverse HA-cyp primers were used for amplification Cytochrome b gene. Mega 10.0 applications were used to construct of the phylogenetic tree. The phylogeographic analysis uses a comparative phylogenetic tree approach with geographical history. The cytochrome b gene showed variations in the nucleotides of O. spilurus from the three islands. Phylogeography of O. spilurus showed that there are two different clades with intraspecific variation according to a geographic area with bootstrap values of 99-100%. One clade contains fish samples from Bangka and Belitung islands, while the other clade comes from Kalimantan island. The formation of the Karimata Strait (between Bangka-Belitung and Kalimantan Islands) and the pattern of paleo rivers are predicted to influence the disparity.

Time for speciation and niche conservatism explain the latitudinal diversity gradient in clupeiform fishes

AbstractAimThe latitudinal diversity gradient of increasing species richness from poles to equator is one of the most striking and pervasive spatial patterns of biodiversity. Climate appears to have been key to the formation of the latitudinal diversity gradient, but the processes through which climate shaped species richness remain unclear. We tested predictions of the time for speciation, carrying capacity and diversification rate latitudinal diversity gradient hypotheses in a trans‐marine/freshwater clade of fishes.LocationGlobal in marine and freshwater environments.TaxonClupeiformes (anchovies, herrings, sardines and relatives).MethodsWe tested predictions of latitudinal diversity gradient hypotheses using a molecular phylogeny, species distribution data and phylogenetic comparative approaches. To test the time for speciation hypothesis, we conducted ancestral state reconstructions to infer the ages of temperate, subtropical and tropical lineages and frequency of evolutionary transitions between climates. We tested the carry capacity hypothesis by characterizing changes in net diversification rates through time. To test the diversification rate hypothesis, we qualitatively compared the diversification rates of temperate, subtropical and tropical lineages and conducted statistical tests for associations between latitude and diversification rates.ResultsWe identified four transitions to temperate climates and two transitions out of temperate climates. We found no differences in diversification rates among temperate and tropical clupeiforms. Net diversification rates remained positive in crown Clupeiformes since their origin ~150 Ma in both tropical and temperate lineages. Climate niche characters exhibited strong phylogenetic signal. All temperate clupeiform lineages arose <50 Ma, after the Early Eocene Climatic Optimum.Main conclusionsOur results support the time for speciation hypothesis, which proposes that climate niche conservatism and fluctuations in the extent of temperate climates limited the time for species to accumulate in temperate climates, resulting in the latitudinal diversity gradient. We found no support for the carrying capacity or diversification rate hypotheses.