Family-level Phylogeny Research Articles

An Opiliones-specific ultraconserved element probe set with a near-complete family-level phylogeny

Sequence capture of ultraconserved elements (UCEs) has transformed molecular systematics across many taxa, with arachnids being no exception. The probe set available for Arachnida has been repeatedly used across multiple arachnid lineages and taxonomic levels, however more specific probe sets for spiders have demonstrated that more UCEs can be recovered with higher probe specificity. In this study, we develop an Opiliones-specific UCE probe set targeting 1915 UCEs using a combination of probes designed from genomes and transcriptomes, as well as the most useful probes from the Arachnida probe set. We demonstrate the effectiveness of this probe set across Opiliones with the most complete family-level phylogeny made to date, including representatives from 61 of 63 currently described families. We also test UCE recovery from historical specimens with degraded DNA, examine population-level data sets, and assess “backwards compatibility” with samples hybridized with the Arachnida probe set. The resulting phylogenies – which include specimens hybridized using both the Opiliones and Arachnida probe sets, historical specimens, and transcriptomes – are largely congruent with previous multi-locus and phylogenomic analyses. The probe set is also “backwards compatible”, increasing the number of loci obtained in samples previously hybridized with the Arachnida probe set, and shows high utility down to shallow population-level divergences. This probe set has the potential to further transform Opiliones molecular systematics, resolving many long-standing taxonomic issues plaguing this lineage.

Jumping continents and major host lineages: phylogeny and diversity of the enigmatic Cryptotropidae (Platyhelminthes: Digenea).

Members of several genera in the digenean superfamily Microphalloidea, namely Renschetrema, Rohdetrema, Cryptotropa, Cephalouterina and Pseudocryptotropa, are characterized by an unusual dorsal position of the genital atrium. In the absence of phylogenetic data, their systematic position has been unstable. In the most recent taxonomic revision of the Microphalloidea, they were housed in three different families. We studied the morphology and obtained DNA sequences of several microphallolideans with a dorsal genital pore, collected from vertebrates in Southeast Asia and South America. We used sequences of the nuclear 28S gene to infer a superfamily-level phylogeny and sequences of the mitochondrial cox1 gene for family-level phylogeny and species-level comparisons. Based on the combination of molecular phylogenetic data and morphological features, we restore the Cryptotropidae and synonymize the Renschetrematidae with the Cryptotropidae. We erect new genera Paracryptotropagen. nov. and Armadoatriumgen. nov., provide amended diagnoses of the Cryptotropidae and Renschetrema and provide keys to the identification of genera within the family. We resurrect the genus Novetrema, previously synonymized with Pseudocryptotropa, and remove Renschetrema indicum from Renschetrema. We describe four new species from lizards in Vietnam and birds in the Philippines and Peru. The unique morphological features and unusually broad host associations and geographical distributions of cryptotropids are discussed.

Comparative mitogenomic analyses provide evolutionary insights into the retrolateral tibial apophysis clade (Araneae: Entelegynae).

The retrolateral tibial apophysis (RTA) clade is the largest spider lineage within Araneae. To better understand the diversity and evolution, we newly determined mitogenomes of ten RTA species from six families and performed a comparative mitogenomics analysis by combining them with 40 sequenced RTA mitogenomes available on GenBank. The ten mitogenomes encoded 37 typical mitochondrial genes and included a large non-coding region (putative control region). Nucleotide composition and codon usage were well conserved within the RTA clade, whereas diversity in sequence length and structural features was observed in control region. A reversal of strand asymmetry in nucleotide composition, i.e., negative AT-skews and positive GC-skews, was observed in each RTA species, likely resulting from mitochondrial gene rearrangements. All protein-coding genes were evolving under purifying selection, except for atp8 whose Ka/Ks was larger than 1, possibly due to positive selection or selection relaxation. Both mutation pressure and natural selection might contribute to codon usage bias of 13 protein-coding genes in the RTA lineage. Phylogenetic analyses based on mitogenomic data recovered a family-level phylogeny within the RTA; {[(Oval calamistrum clade, Dionycha), Marronoid clade], Sparassidae}. This study characterized RTA mitogenomes and provided some new insights into the phylogeny and evolution of the RTA clade.

Phylogenomics and Diversification of the Schistosomatidae Based on Targeted Sequence Capture of Ultra-Conserved Elements.

Schistosomatidae Stiles and Hassall 1898 is a medically significant family of digenetic trematodes (Trematoda: Digenea), members of which infect mammals or birds as definitive hosts and aquatic or amphibious gastropods as intermediate hosts. Currently, there are 17 named genera, for many of which evolutionary interrelationships remain unresolved. The lack of a resolved phylogeny has encumbered our understanding of schistosomatid evolution, specifically patterns of host-use and the role of host-switching in diversification. Here, we used targeted sequence capture of ultra-conserved elements (UCEs) from representatives of 13 of the 17 named genera and 11 undescribed lineages that are presumed to represent either novel genera or species to generate a phylogenomic dataset for the estimation of schistosomatid interrelationships. This study represents the largest phylogenetic effort within the Schistosomatidae in both the number of loci and breadth of taxon sampling. We present a near-comprehensive family-level phylogeny providing resolution to several clades of long-standing uncertainty within Schistosomatidae, including resolution for the placement of the North American mammalian schistosomes, implying a second separate capture of mammalian hosts. Additionally, we present evidence for the placement of Macrobilharzia at the base of the Schistosoma + Bivitellobilharzia radiation. Patterns of definitive and intermediate host use and a strong role for intermediate host-switching are discussed relative to schistosomatid diversification.

Family–level phylogeny of infraorder Systellognatha (Insecta: Plecoptera) inferred from mitochondrial genomes

AbstractSystellognatha is a monophyletic group within stoneflies containing seven families in two superfamilies. However, the family–level phylogeny within Systellognatha is largely unresolved, with some controversial hypotheses generated by molecular data. In this study, we sequenced one complete and three nearly complete mitochondrial genomes (mitogenomes) representing three systellognathan families (including one newly described family), to recover the phylogeny of all seven extant systellognathan families. We tested the effect of heterogeneous sequence divergence within mitogenome sequences on phylogenetic reconstruction. To study incongruent topologies supported by different datasets, we also utilized four–cluster likelihood mapping to elucidate these phylogenetic questions. We found a low degree of compositional heterogeneity in systellognathan mitogenomes. Our results showed that phylogenetic analyses with various datasets under site–homogeneous models failed to recover the Perloidea and Pteronarcyoidea as monophyletic groups. However, Bayesian inference with a site–heterogeneous model (CAT + GTR) recovered Perloidea as monophyletic. Our results confirm a sister–group relationship between Chloroperlidae and Perlodidae. Despite all phylogenetic analyses failed to recover the Pteronarcyoidea as monophyletic, the analyses with site–heterogeneous model based on PCG12, PCG12R and PCGR datasets reconstructed the same relationships among Pteronarcyidae, Peltoperlidae, Styloperlidae and Perloidea. Our study shows the power of the site–heterogeneous models in resolving phylogeny within Systellognatha and provides a new insight into the phylogeny of this group.

Phylogenetic signal of sub‐arctic beetle communities

Postglacial dispersal and colonization processes have shaped community patterns in sub‐Arctic regions such as Churchill, Manitoba, and Canada. This study investigates evolutionary community structure within the beetle (Coleoptera) families of Churchill and tests whether biological traits have played a role in governing colonization patterns from refugial and southerly geographic regions. This study quantifies sub‐Arctic beetle phylogenetic community structure for each family using the net relatedness index (NRI) and nearest taxon index (NTI), calculated using publicly available data from the Barcode of Life Data Systems (BOLD); compares patterns across families with different traits (habitat, diet) using standard statistical analysis (ANOVA) as well as phylogenetic generalized least squares (PGLS) using a family‐level beetle phylogeny obtained from the literature; and compares community structure in Churchill with a region in southern Canada (Guelph, Ontario). These analyses were also repeated at a genus level. The dominant pattern detected in our study was that aquatic families were much better represented in Churchill compared to terrestrial families, when compared against richness sampled from across Canada and Alaska. Individually, most families showed significant phylogenetic clustering in Churchill, likely due to the strong environmental filtering present in Arctic environments. There was no significant difference in phylogenetic structure between Churchill and Guelph but with a trend toward stronger clustering in the North. Fungivores were significantly more overdispersed than other feeding modes, predators were significantly more clustered, and aquatic families showed significantly stronger clustering compared to terrestrial. This study contributes to our understanding of the traits and processes structuring insect biodiversity and macroecological trends in the sub‐Arctic.

Evolutionary Patterns in Sound Production across Fishes

Sound production by fishes has been recognized for millennia, but is typically regarded as comparatively rare and thus yet to be integrated into broader concepts of vertebrate evolution. We map the most comprehensive dataset of sound production yet assembled onto a family-level phylogeny of ray-finned fishes (Actinopterygii), a clade containing more than 34,000 extant species. Family-, rather than species-, level analyses allowed broad investigation of sound production mostly based on illustrations of acoustic recordings and morphological specializations (82%) strongly indicative of sound production along with qualitative descriptions (18%), and a conservative estimate of the distribution and ancestry of a character that is likely more widespread than currently known. Compilation of sonic-related morphological characters shows 60 families exhibiting muscles coupled to swim bladder vibration and 39 families that employ movement of skeletal parts against each other, i.e., stridulation. Eighteen of these families, mostly catfishes (13), include individual species exhibiting both mechanisms. The results show that families with soniferous species contain nearly two-thirds of actinopterygian species, including a clade originating circa 155 Ma, and that sound production has independently evolved approximately 33 times within Actinopterygii. Despite the uncertainties of presence-only data records and incomplete evidence of absence, under-sampling species, and assuming family-level conservation of sound production, sensitivity analyses show that these patterns of shared ancestry are robust. In aggregate, these findings offer a new perspective on the ancestry and convergent evolution of sound production among actinopterygians, a clade representing more than half of extant vertebrate species.

The delayed and geographically heterogeneous diversification of flowering plant families.

The Early Cretaceous (145-100 million years ago (Ma)) witnessed the rise of flowering plants (angiosperms), which ultimately lead to profound changes in terrestrial plant communities. However, palaeobotanical evidence shows that the transition to widespread angiosperm-dominated biomes was delayed until the Palaeocene (66-56 Ma). Important aspects of the timing and geographical setting of angiosperm diversification during this period, and the groups involved, remain uncertain. Here we address these aspects by constructing and dating a new and complete family-level phylogeny, which we integrate with 16 million geographic occurrence records for angiosperms on a global scale. We show substantial time lags (mean, 37-56 Myr) between the origin of families (stem age) and the diversification leading to extant species (crown ages) across the entire angiosperm tree of life. In turn, our results show that families with the shortest lags are overrepresented in temperate and arid biomes compared with tropical biomes. Our results imply that the diversification and ecological expansion of extant angiosperms was geographically heterogeneous and occurred long after most of their phylogenetic diversity originated during the Cretaceous Terrestrial Revolution.

How to build distylous flowers: comparative floral development and evolution of distylous species across the angiosperms.

Distyly, a plant breeding system characterized by two floral morphs that have reciprocal positioning of anthers and stigmas, is known from at least 27 angiosperm families, making it an excellent example of convergent evolution. The various manners in which patterns of floral development produce the distinct anther and stigma heights in each morph remain largely unexplored from developmental and evolutionary perspectives. In 15 species representing at least 12 origins of distyly, heights and lengths of floral organs in each morph throughout development were examined using light microscopy. Patterns of floral organ development were determined and compared among species. Family-level phylogenies of distylous species and relatives were reconstructed, and patterns of ancestral herkogamy were resolved. Differences in floral development between morphs resulted in 12 patterns leading to the anther and stigma positions characterizing distyly. Distylous species evolved from ancestors with different types of herkogamy, with approach herkogamy and lack of herkogamy resolved most frequently. Seven of the 12 patterns of floral development are known from only one species, with three other patterns described among pairs of close relatives. The most common pattern of floral development, described from at least seven genera, involves for anther heights, distinct intermorph growth rates and for stigma heights, growth rates that differ between morphs only during later development. This pattern is common among subclass Lamiidae, suggesting canalized development within the taxon. Among distylous species, the same type of ancestral herkogamy can give rise to different patterns of floral development.

Brassicales phylogeny inferred from 72 plastid genes: A reanalysis of the phylogenetic localization of two paleopolyploid events and origin of novel chemical defenses.

Previous phylogenetic studies employing molecular markers have yielded various insights into the evolutionary history across Brassicales, but many relationships between families remain poorly supported or unresolved. A recent phylotranscriptomic approach utilizing 1155 nuclear markers obtained robust estimates for relationships among 14 of 17 families. Here we report a complete family-level phylogeny estimated using the plastid genome. We conducted phylogenetic analyses on a concatenated data set comprising 44,926 bp from 72 plastid genes for species distributed across all 17 families. Our analysis includes three additional families, Tovariaceae, Salvadoraceae, and Setchellanthaceae, that were omitted in the previous phylotranscriptomic study. Our phylogenetic analyses obtained fully resolved and strongly supported estimates for all nodes across Brassicales. Importantly, these findings are congruent with the topology reported in the phylotranscriptomic study. This consistency suggests that future studies could utilize plastid genomes as markers for resolving relationships within some notoriously difficult clades across Brassicales. We used this new phylogenetic framework to verify the placement of the At-α event near the origin of Brassicaceae, with median date estimates of 31.8 to 42.8 million years ago and restrict the At-β event to one of two nodes with median date estimates between 85 to 92.2 million years ago. These events ultimately gave rise to novel chemical defenses and are associated with subsequent shifts in net diversification rates. We anticipate that these findings will aid future comparative evolutionary studies across Brassicales, including selecting candidates for whole-genome sequencing projects.

Shallow size-density relations within mammal clades suggest greater intra-guild ecological impact of large-bodied species.

Population densities of species have a predictable relationship with their body mass on a global scale. This relationship is known as the size-density relationship (SDR). The relationship was originally found to be directly opposite of metabolic rate scaling, which led to the hypothesis of energetic equivalence. However, recent studies have suggested that the SDR varies between clades. Specifically, the SDR for certain mammal clades has been found to be less negative than the relationship across all mammals. The aim of the present study is to estimate phylogenetic variation in the scaling relationship, using a data-driven identification of natural phylogenetic substructure in the body size-density relation, and discuss its potential drivers. The classic model is often used to estimate natural population densities, and a further, practical aim is to improve it by incorporating variability among phylogenetic groups. We expand the model for the SDR relation of mammals to include clade-specific variation. We used a dataset with population and body mass estimates of 924 terrestrial mammal species, covering 97 families, and applied an algorithm identifying group-specific changes in the relationship across a family-level phylogeny. We show increased performance in species density estimation is achieved by incorporating clade-specific changes in the relationship compared to the classic model (increasing r2 from .56 to .74 and ΔAICc =466). While the global SDR across clades was confirmed to be similar to previous findings (r=-.74), the relationship within all sub-clades was less negative than the overall trend. Our results show that data-driven identification of phylogenetic substructure in the size-density relation substantially improves predictive accuracy of the model. The less negative relationship within clades compared to the overall trend and compared to within clade metabolic scaling suggest that the energetic equivalence rule does not hold. This relationship shows that large species within clades use proportionally more energy than smaller species. Therefore, our results are consistent with a greater intra-guild ecological impact of large-bodied species via partial monopolisation of resources by the largest species of a given guild, and hence size-asymmetric intra-guild competition.

Natural Constraints to Species Diversification.

Identifying modes of species diversification is fundamental to our understanding of how biodiversity changes over evolutionary time. Diversification modes are captured in species phylogenies, but characterizing the landscape of diversification has been limited by the analytical tools available for directly comparing phylogenetic trees of groups of organisms. Here, we use a novel, non-parametric approach and 214 family-level phylogenies of vertebrates representing over 500 million years of evolution to identify major diversification modes, to characterize phylogenetic space, and to evaluate the bounds and central tendencies of species diversification. We identify five principal patterns of diversification to which all vertebrate families hold. These patterns, mapped onto multidimensional space, constitute a phylogenetic space with distinct properties. Firstly, phylogenetic space occupies only a portion of all possible tree space, showing family-level phylogenies to be constrained to a limited range of diversification patterns. Secondly, the geometry of phylogenetic space is delimited by quantifiable trade-offs in tree size and the heterogeneity and stem-to-tip distribution of branching events. These trade-offs are indicative of the instability of certain diversification patterns and effectively bound speciation rates (for successful clades) within upper and lower limits. Finally, both the constrained range and geometry of phylogenetic space are established by the differential effects of macroevolutionary processes on patterns of diversification. Given these properties, we show that the average path through phylogenetic space over evolutionary time traverses several diversification stages, each of which is defined by a different principal pattern of diversification and directed by a different macroevolutionary process. The identification of universal patterns and natural constraints to diversification provides a foundation for understanding the deep-time evolution of biodiversity.

Community Phylogenetics: Assessing Tree Reconstruction Methods and the Utility of DNA Barcodes.

Studies examining phylogenetic community structure have become increasingly prevalent, yet little attention has been given to the influence of the input phylogeny on metrics that describe phylogenetic patterns of co-occurrence. Here, we examine the influence of branch length, tree reconstruction method, and amount of sequence data on measures of phylogenetic community structure, as well as the phylogenetic signal (Pagel’s λ) in morphological traits, using Trichoptera larval communities from Churchill, Manitoba, Canada. We find that model-based tree reconstruction methods and the use of a backbone family-level phylogeny improve estimations of phylogenetic community structure. In addition, trees built using the barcode region of cytochrome c oxidase subunit I (COI) alone accurately predict metrics of phylogenetic community structure obtained from a multi-gene phylogeny. Input tree did not alter overall conclusions drawn for phylogenetic signal, as significant phylogenetic structure was detected in two body size traits across input trees. As the discipline of community phylogenetics continues to expand, it is important to investigate the best approaches to accurately estimate patterns. Our results suggest that emerging large datasets of DNA barcode sequences provide a vast resource for studying the structure of biological communities.

Re-evaluating the phylogeny of Sipuncula through transcriptomics

Sipunculans (also known as peanut worms) are an ancient group of exclusively marine worms with a global distribution and a fossil record that dates back to the Early Cambrian. The systematics of sipunculans, now considered a distinct subclade of Annelida, has been studied for decades using morphological and molecular characters, and has reached the limits of Sanger-based approaches. Here, we reevaluate their family-level phylogeny by comparative transcriptomic analysis of eight species representing all known families within Sipuncula. Two data matrices with alternative gene occupancy levels (large matrix with 675 genes and 62% missing data; reduced matrix with 141 genes and 23% missing data) were analysed using concatenation and gene-tree methods, yielding congruent results and resolving each internal node with maximum support. We thus corroborate prior phylogenetic work based on molecular data, resolve outstanding issues with respect to the familial relationships of Aspidosiphonidae, Antillesomatidae and Phascolosomatidae, and highlight the next area of focus for sipunculan systematics.

Is inaccurate mimicry ancestral to accurate in myrmecomorphic spiders (Araneae)?

Batesian mimicry, in which a palatable organism resembles an unpalatable model, is widespread among taxa. Batesian mimics can be classified based on their level of accuracy (inaccurate or accurate). Using data on defensive strategies in more than 1000 species of spiders I investigated whether inaccurate myrmecomorphy is ancestral to accurate myrmecomorphy. I classified 233 myrmecomorphic species into four accuracy levels based on morphology, from poor inaccurate mimics to very accurate ones. I found that myrmecomorphy has evolved independently in 16 families and 85 genera. On the family-level phylogeny, the occurrence of myrmecomorphy is confined mainly to families branching later on the tree, from the RTA clade. On the generic-level phylogenies in Corinnidae and Salticidae, myrmecomorphy is not only of derived origin. Estimated ancestral state was non-mimetic in Salticidae and poor inaccurate myrmecomorphy in Corinnidae. Thus, inaccurate myrmecomorphic spider mimics seem rather ancestral to accurate but additional analysis on species-level phylogenies is needed to support this conclusion. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113, 97–111.

A molecular phylogeny of Pacific honeyeaters (Aves: Meliphagidae) reveals extensive paraphyly and an isolated Polynesian radiation

We investigated the molecular phylogenetic placement of 14 species of Pacific island honeyeaters (Aves: Meliphagidae) in the broader context of an existing family-level phylogeny. We examined the evolutionary history of Pacific honeyeater lineages to assess the accuracy of current taxonomies and to evaluate their biogeographic history. We compare these biogeographic patterns to other Pacific birds to identify emergent patterns across lineages. We found strong support for a previously unknown endemic radiation in central Polynesia, which comprises five genera: Meliarchus, Guadalcanaria, Gymnomyza, Xanthotis, and Foulehaio. Conversely, other Pacific lineages were found to be strongly allied with continental radiations (e.g., Philemon eichhorni, P. cockerelli, and Lichmera incana). Our results necessitated taxonomic changes, both at the generic level (e.g., Xanthotis, Melidectes/Vosea, and Glycifohia/Gliciphila) and regarding species limits within polytypic species. Here, we discuss species limits in Foulehaio and Gymnomyza and recommend elevating three nominal subspecies of Foulehaio to species status, each of which forms well-differentiated clades.

Resolving deep lineage divergences in core corvoid passerine birds supports a proto-Papuan island origin

It is well established that the global expansion of songbirds (Oscines) originated in East Gondwana (present day Australo-Papua), and it has been postulated that one of the main constituent groups, the “core Corvoidea”, with more than 750 species, originated in the first islands that emerged where New Guinea is now located. However, several polytomous relationships remained within the clade, obstructing detailed biogeographical interpretations. This study presents a well-resolved family-level phylogeny, based on a dataset of 22 nuclear loci and using a suite of partitioning schemes and Maximum Likelihood and Bayesian inference methods. Resolving the relationships within the core Corvoidea provides evidence for three well-supported main clades, which are in turn sister to the New Zealand genus Mohoua. Some monotypic lineages, which have previously been considered Incertae sedis, are also placed in a phylogenetic context. The well-resolved phylogeny provides a robust framework for biogeographical analyses, and provides further support for the hypothesis that core corvoids originated in the proto-Papuan island region that emerged north of Australia in the late Oligocene/early Miocene. Thus, the core Corvoidea appear to represent a true island radiation, which successfully colonized all continents except Antarctica.

Into the deep: A phylogenetic approach to the bivalve subclass Protobranchia

A molecular phylogeny of Protobranchia, the subclass of bivalve mollusks sister to the remaining Bivalvia, has long proven elusive, because many constituent lineages are deep-sea endemics, which creates methodological challenges for collecting and preserving genetic material. We obtained 74 representatives of all 12 extant protobranch families and investigated the internal phylogeny of this group using sequence data from five molecular loci (16S rRNA, 18S rRNA, 28S rRNA, cytochrome c oxidase subunit I, and histone H3). Model-based and dynamic homology parsimony approaches to phylogenetic reconstruction unanimously supported four major clades of Protobranchia, irrespective of treatment of hypervariable regions in the nuclear ribosomal genes 18S rRNA and 28S rRNA. These four clades correspond to the superfamilies Nuculoidea (excluding Sareptidae), Nuculanoidea (including Sareptidae), Solemyoidea, and Manzanelloidea. Salient aspects of the phylogeny include (1) support for the placement of the family Sareptidae with Nuculanoidea; (2) the non-monophyly of the order Solemyida (Solemyidae+Nucinellidae); (3) and the non-monophyly of most nuculoid and nuculanoid genera and families. In light of this first family-level phylogeny of Protobranchia, we present a revised classification of the group. Estimation of divergence times in concert with analyses of diversification rates demonstrate the signature of the end-Permian mass extinction in the phylogeny of extant protobranchs.

Morphological traits as predictors of diet and microhabitat use in a diverse beetle assemblage

We explored how morphological traits can complement phylogenetic information to extend our predictions of the ecology of a diverse beetle assemblage. We analysed ten morphological traits from an assemblage of 239 species from 35 families, and identified three axes of morphological variation that were independent of body length: (1) relative robustness; (2) relative appendage length; and (3) relative abdomen length. The trait associations defining these axes of morphological variation did not change after adjusting for family-level phylogeny. We detected significant differences in morphological variation across the beetle assemblage according to diet and microhabitat use, and these patterns were only partially influenced by family membership. Further analysis within dominant families showed that species of Carabidae, Curculionidae, Scarabaeidae and Staphylinidae had greater body length in open versus tree litter microhabitat, and species of Carabidae and Curculionidae had greater relative robustness, but shorter relative appendage length, in open versus tree litter microhabitat. Although it is clear that family-level phylogeny and morphology share some explanatory power for predicting the diet and microhabitat use by beetles, we demonstrate that body length, robustness and appendage length are correlated significantly with microhabitat use when comparing members of the same family. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102, 301–310.

The origins of species richness in the Hymenoptera: insights from a family-level supertree.

BackgroundThe order Hymenoptera (bees, ants, wasps, sawflies) contains about eight percent of all described species, but no analytical studies have addressed the origins of this richness at family-level or above. To investigate which major subtaxa experienced significant shifts in diversification, we assembled a family-level phylogeny of the Hymenoptera using supertree methods. We used sister-group species-richness comparisons to infer the phylogenetic position of shifts in diversification.ResultsThe supertrees most supported by the underlying input trees are produced using matrix representation with compatibility (MRC) (from an all-in and a compartmentalised analysis). Whilst relationships at the tips of the tree tend to be well supported, those along the backbone of the tree (e.g. between Parasitica superfamilies) are generally not. Ten significant shifts in diversification (six positive and four negative) are found common to both MRC supertrees. The Apocrita (wasps, ants, bees) experienced a positive shift at their origin accounting for approximately 4,000 species. Within Apocrita other positive shifts include the Vespoidea (vespoid wasps/ants containing 24,000 spp.), Anthophila + Sphecidae (bees/thread-waisted wasps; 22,000 spp.), Bethylidae + Chrysididae (bethylid/cuckoo wasps; 5,200 spp.), Dryinidae (dryinid wasps; 1,100 spp.), and Proctotrupidae (proctotrupid wasps; 310 spp.). Four relatively species-poor families (Stenotritidae, Anaxyelidae, Blasticotomidae, Xyelidae) have undergone negative shifts. There are some two-way shifts in diversification where sister taxa have undergone shifts in opposite directions.ConclusionsOur results suggest that numerous phylogenetically distinctive radiations contribute to the richness of large clades. They also suggest that evolutionary events restricting the subsequent richness of large clades are common. Problematic phylogenetic issues in the Hymenoptera are identified, relating especially to superfamily validity (e.g. "Proctotrupoidea", "Mymarommatoidea"), and deeper apocritan relationships. Our results should stimulate new functional studies on the causes of the diversification shifts we have identified. Possible drivers highlighted for specific adaptive radiations include key anatomical innovations, the exploitation of rich host groups, and associations with angiosperms. Low richness may have evolved as a result of geographical isolation, specialised ecological niches, and habitat loss or competition.