Discrete Character Research Articles

Rapid Initial Morphospace Expansion and Delayed Morphological Disparity Peak in the First 100 Million Years of the Archosauromorph Evolutionary Radiation

Adaptive radiations have played a major role in generating modern and deep-time biodiversity. The Triassic radiation of the Archosauromorpha was one of the most spectacular vertebrate radiations, giving rise to many highly ecomorphologically varied lineages—including the dinosaurs, pterosaurs, and stem-crocodylians—that dominated the larger-bodied land fauna for the following 150 Ma, and ultimately gave rise to today’s > 10,000 species of birds and crocodylians. This radiation provides an outstanding testbed for hypotheses relating to adaptive radiations more broadly. Recent studies have started to characterize the tempo and mode of the archosauromorph early adaptive radiation, indicating very high initial rates of evolution, non-competitive niche-filling processes, and previously unrecognized morphological disparity even among non-crown taxa. However, these analyses rested primarily either on discrete characters or on geometric morphometrics of the cranium only, or even failed to fully include phylogenetic information. Here we expand previous 2D geometric morphometric cranial datasets to include new taxa and reconstructions, and create an analogous dataset of the pelvis, thereby allowing comparison of anatomical regions and the transition from “sprawling” to “upright” posture to be examined. We estimated morphological disparity and evolutionary rates through time. All sampled clades showed a delayed disparity peak for sum of variances and average nearest neighbor distances in both the cranium and pelvis, with disparity likely not saturated by the end of the studied time span (Late Jurassic); this contrasts with smaller radiations, but lends weight to similar results for large, ecomorphologically-varied groups. We find lower variations in pelvic than cranial disparity among Triassic-Jurassic archosaurs, which may be related to greater morphofunctional constraints on the pelvis. Contrasting with some previous work, but also confirming some previous findings during adaptive radiations, we find relatively widespread evidence of correlation between sampled diversity and disparity, especially at the largest phylogenetic scales and using average displacement rather than sum of variances as disparity metric; this also demonstrates the importance of comparing disparity metrics, and the importance of phylogenetic scale. Stem and crown archosauromorphs show a morphological diversification of both the cranium and pelvis with higher initial rates (Permian–Middle Triassic and at the base of major clades) followed by lower rates once diversification into niches has occurred (Late Triassic–Jurassic), indicating an “early burst” patternsensu lato. Our results provide a more detailed and comprehensive picture of the early archosauromorph radiation and have significant bearing on the understanding of deep-time adaptive radiations more broadly, indicating widespread patterns of delayed disparity peaks, initial correlation of diversity and disparity, and evolutionary early bursts.

Phylogeny, disparity and mass extinction response in the trilobite order Harpetida

AbstractThe trilobite order Harpetida has long been easily recognized but many unanswered evolutionary questions about the group remain. This work explores the phylogenetic relationships within Harpetida and studies the harpetid response to the Late Ordovician mass extinction to better understand the relationship between extinction events and disparity. A discrete morphological character matrix was assembled from published descriptions and refined through first‐hand observations. This matrix is the first attempt of its kind to characterize the overall morphology of Harpetida, rather than focusing on individual harpetid genera. Phylogenetic analyses under both maximum parsimony and Bayesian inference optimality criteria retrieve tree topologies that support harpetid monophyly but throw doubt onto previous hypotheses of the internal relationships of the order. Harpetid disparity proves remarkably stable over time. A modest peak in the Ordovician is followed by a slow decline throughout the Silurian and Devonian. After the Ordovician period, harpetids demonstrate little or no ability to colonize new areas of morphospace. This may represent a fundamental failure to recover, in which the lasting impacts of Late Ordovician mass extinction continue to suppress morphological innovation. These findings demonstrate that mass extinction events may have complex impacts that play out over many millions of years.

Morphometrics and Redescription of Aphyllon fasciculatum and Aphyllon franciscanum, Two Widespread but Previously Conflated Species in Western North America

Abstract— We continue the taxonomic reevaluation of Aphyllon sect. Aphyllon by describing a widespread species throughout western North America previously recognized within a polyphyletic A. fasciculatum. To support our description and revised key, we analyzed fifteen continuous and discrete characters sampled from 186 herbarium specimens and iNaturalist observations representing the geographic and host ranges. Principal component and multiple correspondence analyses reveal clear variation in floral characters. Discriminant analyses show that three aspects of floral color, corolla lobe tip shape, the calyx cup to calyx lobe ratio, calyx lobe length, and degree of bend in the corolla tube are useful distinguishing features, but not diagnostic in every case.

The phylogenetic relationships of geoemydid turtles from the Eocene Messel Pit Quarry: a first assessment using methods for continuous and discrete characters.

The geoemydid turtles of the Eocoene Messel Pit Quarry of Hesse, Germany, are part of a rich Western European fossil record of testudinoids. Originally referred to as “Ocadia” kehreri and “Ocadia” messeliana, their systematic relationships remain unclear. A previous study proposed that a majority of the Western European geoemydids, including the Messel geoemydids, are closely related to the Recent European representatives of the clade Mauremys. Another study hypothesised that the Western European geoemydid fauna is more phylogenetically diverse, and that the Messel geoemydids are closely related to the East Asian turtles Orlitia and Malayemys. Here we present the first quantitative analyses to date that investigate this question. We use continuous characters in the form of ratios to estimate the placement of the Messel geoemydids in a reference tree that was estimated from molecular data. We explore the placement error obtained from that data with maximum likelihood and Bayesian methods, as well as linear parsimony in combination with discrete characters. We find good overall performance with Bayesian and parsimony analyses. Parsimony performs even better when we also incorporated discrete characters. Yet, we cannot pin down the position of the Messel geoemydids with high confidence. Depending on how intraspecific variation of the ratio characters is treated, parsimony favours a placement of the Messel fossils sister to Orlitia borneensis or sister to Geoemyda spengleri, with weak bootstrap support. The latter placement is suspect because G. spengleri is a phylogenetically problematic species with molecular and morphological data. There is even less support for placements within the Mauremys clade.

Multiple paths to morphological diversification during the origin of amniotes.

Early terrestrial vertebrates (amniotes) provide a classic example of diversification following adaptive zone invasion. The initial terrestrialization of vertebrates was closely followed by dietary diversification, as evidenced by a proliferation of craniomandibular and dental adaptations. However, morphological evolution of early amniotes has received limited study, in analyses with restricted taxonomic scope, leaving substantial questions about the dynamics of this important terrestrial radiation. We use novel analyses of discrete characters to quantify variation in evolutionary rates and constraints during diversification of the amniote feeding apparatus. We find evidence for an early burst, comprising high rates of anatomical change that decelerated through time, giving way to a background of saturated morphological evolution. Subsequent expansions of phenotypic diversity were not associated with increased evolutionary rates. Instead, variation in the mode of evolution became important, with groups representing independent origins of herbivory evolving distinctive, group-specific morphologies and thereby exploring novel character-state spaces. Our findings indicate the importance of plant-animal interactions in structuring the earliest radiation of amniotes and demonstrate the importance of variation in modes of phenotypic divergence during a major evolutionary radiation.

Diversity and evolution of the extraordinary vocal sacs of casque-headed treefrogs (Anura: Hylidae)

Abstract Vocal sacs are among the most conspicuous features of anurans and are particularly striking in casque-headed treefrogs (Hylidae: Hylinae: Lophyohylini) with their wide array of morphologies. In this paper, we assessed the anatomy of vocal sacs in representatives of the Lophyohylini, described eight discrete characters and studied their evolution. We inferred that dorsolateral projections of the vocal sacs were already present during the early evolution of the tribe. Subsequently, they reached surprising volumes in some species, whereas in others they were notably reduced. We inferred between nine and 11 independent events of reduction of the size and lateral projections of the vocal sac, showing unprecedented levels of plasticity for the structure. Moreover, these events were strongly correlated with the colonization of phytotelmata as breeding sites, probably due to their confined space which hampers the inflation of large vocal sacs. Finally, we discuss the evolution of paired lateral vocal sacs in different groups of anurans, and the extent to which the paired and dorsally-projecting lobes of most Trachycephalus differ from those of distantly related taxa. Our findings highlight how variation in internal structure affects the shape of the inflated vocal sac and provides a framework applicable across the Anura.

(2, 2) Scattering and the celestial torus

Analytic continuation from Minkowski space to (2, 2) split signature spacetime has proven to be a powerful tool for the study of scattering amplitudes. Here we show that, under this continuation, null infinity becomes the product of a null interval with a celestial torus (replacing the celestial sphere) and has only one connected component. Spacelike and timelike infinity are time-periodic quotients of AdS3. These three components of infinity combine to an S3 represented as a toric fibration over the interval. Privileged scattering states of scalars organize into SL(2, ℝ)L×SL(2, ℝ)R conformal primary wave functions and their descendants with real integral or half-integral conformal weights, giving the normally continuous scattering problem a discrete character.

The BNSR-invariants of the Stein group 𝐹2,3

Abstract The Stein group F 2 , 3 F_{2,3} is the group of orientation-preserving piecewise linear homeomorphisms of the unit interval with slopes of the form 2 p ⁢ 3 q 2^{p}3^{q} ( p , q ∈ Z p,q\in\mathbb{Z} ) and breakpoints in Z ⁢ [ 1 6 ] \mathbb{Z}[\frac{1}{6}] . This is a natural relative of Thompson’s group 𝐹. In this paper, we compute the Bieri–Neumann–Strebel–Renz (BNSR) invariants Σ m ⁢ ( F 2 , 3 ) \Sigma^{m}(F_{2,3}) of the Stein group for all m ∈ N m\in\mathbb{N} . A consequence of our computation is that (as with 𝐹) every finitely presented normal subgroup of F 2 , 3 F_{2,3} is of type F ∞ \operatorname{F}_{\infty} . Another, more surprising, consequence is that (unlike 𝐹) the kernel of any map F 2 , 3 → Z F_{2,3}\to\mathbb{Z} is of type F ∞ \operatorname{F}_{\infty} , even though there exist maps F 2 , 3 → Z 2 F_{2,3}\to\mathbb{Z}^{2} whose kernels are not even finitely generated. In terms of BNSR-invariants, this means that every discrete character lies in Σ ∞ ⁢ ( F 2 , 3 ) \Sigma^{\infty}(F_{2,3}) , but there exist (non-discrete) characters that do not even lie in Σ 1 ⁢ ( F 2 , 3 ) \Sigma^{1}(F_{2,3}) . To the best of our knowledge, F 2 , 3 F_{2,3} is the first group whose BNSR-invariants are known exhibiting these properties.

SOME THOUGHTS ON INTERSPECIFIC MANDIBULAR MORPHOLOGY IN FOSSIL AND MODERN ALLIGATOR

Alligator is diagnosed in the fossil record using discrete morphological characters. These characters are used with systematic analyses to determine hypothetical phylogenetic relationships. Examined here are two such characters for applicability in fossil species determination and relationship. One is the curvature of the dentary between the fourth and tenth alveoli (observed in several taxa), while the other is a further investigation into the anterior extent of the splenial (in modern Alligator mississippiensis and the early Pleistocene A. hailensis). In a small sample size, the curvature of the dentary exhibits wide variability in character states both intraspecifically and interspecifically, bringing into question its utility in fossil crocodylian systematic applications. The anatomy of the anterior extent of the splenial is phylogenetically informative, and a new method for inferring its presence as a basal or derived state (as a scar on the dentary) is described.

Integrative taxonomy resolves species identities within the Macrobiotus pallarii complex (Eutardigrada: Macrobiotidae)

The taxonomy of many groups of meiofauna is challenging due to their low number of diagnostic morphological characters and their small body size. Therefore, with the advent of molecular techniques that provide a new source of traits, many cryptic species have started to be discovered. Tardigrades are not an exception, and many once thought to be cosmopolitan taxa are being found to be complexes of phenotypically similar species. Macrobiotus pallarii Maucci, 1954 was originally described in South Italy and has been subsequently recorded in Europe, America, and Asia. This allegedly wide geographic range suggests that multiple species may be hidden under this name. Moreover, recently, genetic evidence to support this was put forward, and the Macrobiotus pallarii complex has been proposed to accommodate putative species related to M. pallarii. Here, we describe three new pseudocryptic species based on populations that would have been all classified as Macrobiotus pallarii if molecular methods were not employed. Using an integrative taxonomy approach, we analyzed animals and eggs from the topotypic population of Macrobiotus pallarii, together with four other populations of the complex. We recovered four distinct phylogenetic lineages that, despite the overlap of morphometric traits, can be separated phenotypically by subtle but discrete morphological characters. One lineage corresponds to Macrobiotus pallarii, whereas the other three are newly described as Macrobiotus margoae Stec, Vecchi & Bartels, sp. nov. from the USA, Macrobiotus ripperi Stec, Vecchi & Michalczyk, sp. nov. from Poland and Finland, and Macrobiotus pseudopallarii Stec, Vecchi & Michalczyk, sp. nov. from Montenegro. To facilitate species identification, we provide a dichotomous key for species of the M. pallarii complex. Delimitation of these pseudocryptic taxa highlights the need for an integrative approach to uncover the phylum’s diversity in full.

Evolution of flower morphology and a natural re‐arrangement of Calyceraceae

AbstractCalyceraceae is a small South American family sister to Asteraceae. Previous phylogenetic analyses revealed that Calyceraceae genera are para‐ or polyphyletic. Fruit and inflorescence characters used historically as diagnostic in Calyceraceae taxonomy have proven unreliable in defining natural groups. Surprisingly, flower morphology does not appear in diagnoses of Calyceraceae genera, and its diversity and evolution has remained unstudied before now. Based on phylogenetic analyses using six molecular markers and an almost complete sampling of the family (43 of 46 species), we propose a taxonomic re‐arrangement of Calyceraceae. In the search for diagnostic morphological characters, we studied flower structure diversity and optimized 12 discrete characters by maximum parsimony and maximum likelihood to reconstruct floral evolution in the family. Calyceraceae species are arranged in eight genera (three monotypic, five with 6 to 13 species). Two genera are new (Anachoretes, Asynthema), one is reinstated (Leucocera), with the remaining five emended (Acicarpha, Boopis, Calycera, Gamocarpha, Moschopsis). Flower morphology is notably diverse but provides only a few diagnostic features. Many floral attributes likely reflect selection for pollination or fruit dispersal rather than phylogenetic affinity. In the context of this taxonomic revision, we include a key to genera, new and emended diagnoses, and new combinations.

Phylogeny of true ladybird beetles (Coccinellidae: Coccinellini) reveals pervasive convergent evolution and a rapid Cenozoic radiation

AbstractCoccinellini, the ‘true ladybirds’, is a large, cosmopolitan tribe of lady beetles with immense variability in colour and form that include many commonly recognized species and major biocontrol agents, as well as some ubiquitous invasive species. In this study, we performed phylogenetic analyses of molecular and morphological data. For the molecular analysis, we gathered DNA sequence data from 164 species in 57 genera (and outgroups) from five nuclear genes [CAD (two fragments), TOPO, WGL, 3059fin] and one mitochondrial (COI) gene. The morphological analyses comprised 301 terminals and 61 discrete characters that were analysed by parsimony (MP). Our phylogenetic analyses indicate a monophyletic Coccinellini (former Coccinellinae) containing previously recognized tribes [Coccinellini, Discotomini, Halyziini, Singhikaliini and Tytthaspidini (incl. Bulaeini)]. Coccinellini is further divided into four main clades as follows: the Coccinella group (Clade A; incl. former Tytthaspidini), the Cycloneda group (Clade B), the Hippodamia group (Clade C) and the Synonycha group (Clade D; incl. former Halyziini, Singhikaliini and Discotomini). The phylogenetic relationships between them are: (A, (B, (C, D))). Our analyses also indicate that several currently recognized genera of Coccinellini are para‐ or polyphyletic. Based on our phylogenetic results, new generic groups are defined for Coccinellini, and new generic amendments are proposed as follows: Micraspis Chevrolat sensu n. (Oriental and Australian species), Microcaria Crotch sensu n. (=Bothrocalvia Crotch) (incl. Coelophora Mulsant species) and Phrynocaria Timberlake sensu n. (incl. Coelophora species). Further possible synonyms are discussed, for example, Coccinella Linnaeus (incl. Lioadalia Crotch), Psyllobora Chevrolat (including Oxytella Weise) and Xanthadalia Crotch (incl. Declivitata Fürsch and the African species of Micraspis). The large Clade D, the Synonycha group, with 26 predominant Old World genera, contains the two notable, large and non monophyletic genera Calvia Mulsant and Coelophora that confound the relationships throughout the clade. Additionally, we performed ancestral state reconstruction of selected morphological characters traditionally used in the classification of this group, revealing their homoplastic nature. We also found that the traditional synapomorphy of the tribe, that is, female colleterial glands, closely resemble structures used in symbiotic relationships in other beetles. We used the results of our phylogenetic inferences to estimate divergence times of the tribe and major clades of Coccinellini.

Evolution of the strikingly diverse submandibular muscles in Anura.

The most ventral muscles of the head (the mm. submentalis, intermandibularis, and interhyoideus) provide support to the gular region and lift the buccal floor during ventilation and feeding. These muscles show limited variation in most gnathostomes, but in Anura they exhibit a surprising diversity. The few studies that have explored this character system highlighted its potential as a source of phylogenetic information. In this paper we explored the diversity of this character system studying specimens of 567 anuran species and reviewing published data to cover a total of 1321 species, belonging to 53 of the 54 currently recognized anuran families, as well as caudates and caecilians. We defined 27 discrete characters including the number of muscle bellies, supplementary layers, hypertrophy and diversity of elastic fibres, and pigmentation, among others, and optimized them on a comprehensive phylogenetic hypothesis. We recognized 223 unambiguously optimized synapomorphies for numerous clades on different scales, including three for Anura and many for suprafamiliar clades with poor phenotypic support. Finally, we discussed the evolution of this highly diverse character system, including homology, development, and its functional role in vocalization and feeding. Interestingly, the striking levels of variation in some structures contrast with the amount of phylogenetic inertia, allowing us to recognize several general patterns. Supplementary elements of the m. intermandibularis evolved first as broad layers occuring in more than half of extant anuran species and then concentrated forming discreet bellies in several clades. The anterior portion of the gular region is not sexually dimorphic, and is likely related to ventilation and tongue protraction. Conversely, the diversity of the m. interhyoideus is strongly linked to vocal sacs, which are present only in adult males, suggesting the presence of two independent modules.

The patterns and modes of the evolution of disparity in Mesozoic birds.

The origin of birds from non-avian theropod dinosaurs is one of the greatest transitions in evolution. Shortly after diverging from other theropods in the Late Jurassic, Mesozoic birds diversified into two major clades-the Enantiornithes and Ornithuromorpha-acquiring many features previously considered unique to the crown group along the way. Here, we present a comparative phylogenetic study of the patterns and modes of Mesozoic bird skeletal morphology and limb proportions. Our results show that the major Mesozoic avian groups are distinctive in discrete character space, but constrained in a morphospace defined by limb proportions. The Enantiornithines, despite being the most speciose group of Mesozoic birds, are much less morphologically disparate than their sister clade, the Ornithuromorpha-the clade that gave rise to living birds, showing disparity and diversity were decoupled in avian history. This relatively low disparity suggests that diversification of enantiornithines was characterized in exhausting fine morphologies, whereas ornithuromorphs continuously explored a broader array of morphologies and ecological opportunities. We suggest this clade-specific evolutionary versatility contributed to their sole survival of the end-Cretaceous mass extinction.

Infra-specific variation of Acer cappadocicum (Sapindaceae): morphological and molecular approaches

Besides Quercus, Acer L. genus is the largest tree genus in the Northern Hemisphere and consists of about 124–156 species. High morphological and molecular variability has led to infra-specific complexity within most species of this genus. In Iran, the Acer cappadocicum subsp. cappadocicum Gled. is distributed in eastern to western parts of the Hyrcanian forests. In this study, three leaf-morphotypes in A. cappadocicum Gled. were identified during sampling in a forest located in the Hyrcanian region, leading to obscuring taxonomic situations of this taxon. Against this backdrop, the present study was performed to clarify intraspecific situation of this valuable tree using both macro-/micromorphological and molecular approaches. Forty-five plant samples of A. cappadocicum were randomly selected within nine geographical regions of four provinces in Hyrcanian forests of Iran. Internal transcribed spacer (ITS) (for eight individuals) and inter-simple sequence repeat (ISSR) molecular markers (for 45 individuals) were used along with the epidermal scanning electron micrograph (SEM) of the leaf epidermis. Considering the obtained results, acceptance of the expected lower taxa in A. cappadocicum ssp. cappadocicum in north of Iran could not be supported by molecular evidence. Results of the study indicated no correlation between variations in leaf-morphology, ISSRs, and ITS. However, there are two main clusters in ITS tree without discrete morphological characters showing differentiation and variability in ITS region within this species. This subspecies was compared also with other subspecies.

Results from the Ice Thickness Models Intercomparison eXperiment Phase 2 (ITMIX2)

Knowing the ice thickness distribution of a glacier is of fundamental importance for a number of applications, ranging from the planning of glaciological fieldwork to the assessments of future sea-level change. Across spatial scales, however, this knowledge is limited by the paucity and discrete character of available thickness observations. To obtain a spatially coherent distribution of the glacier ice thickness, interpolation or numerical models have to be used. Whilst the first phase of the Ice Thickness Models Intercomparison eXperiment (ITMIX) focused on approaches that estimate such spatial information from characteristics of the glacier surface alone, ITMIX2 sought insights for the capability of the models to extract information from a limited number of thickness observations. The analyses were designed around 23 test cases comprising both real-world and synthetic glaciers, with each test case comprising a set of 16 different experiments mimicking possible scenarios of data availability. A total of 13 models participated in the experiments. The results show that the inter-model variability in the calculated local thickness is high, and that for unmeasured locations, deviations of 16% of the mean glacier thickness are typical (median estimate, three-quarters of the deviations within 37% of the mean glacier thickness). This notwithstanding, limited sets of ice thickness observations are shown to be effective in constraining the mean glacier thickness, demonstrating the value of even partial surveys. Whilst the results are only weakly affected by the spatial distribution of the observations, surveys that preferentially sample the lowest glacier elevations are found to cause a systematic underestimation of the thickness in several models. Conversely, a preferential sampling of the thickest glacier parts proves effective in reducing the deviations. The response to the availability of ice thickness observations is characteristic to each approach and varies across models. On average across models, the deviation between modeled and observed thickness increase by 8.5% of the mean ice thickness every time the distance to the closest observation increases by a factor of 10. No single best model emerges from the analyses, confirming the added value of using model ensembles.

Planning for sustainable urban mobility: Demand estimation of on-street vertical walking facilities

The improvement of pedestrian mobility through the use of on-street vertical facilities, i.e. escalators, moving sidewalks, lifts, funiculars and gondola lifts, is becoming increasingly important in many cities with steep slopes that hinder walkers’ transit. However, studies on the acceptance, profitability and demand for these facilities are still scarce. The present paper proposes an assessment methodology to evaluate the demand for vertical pedestrian routes and facilities based on diverse accessibility indicators. This methodology was applied to the city of Santander (Spain), where flow measurements and surveys among users of existing vertical walking systems were carried out. These data were modeled using Poisson and Binomial Negative regressions, considering the truncated and discrete character of the dependent variable. The results obtained confirm that vertical walking facilities are highly valued by users. Contrary to expectations, accessibility from these facilities to public services or commercial areas was not relevant to estimate the demand of the existing routes and only accessibility to population proved to be significant. Therefore, it is advisable to install this type of walking facilities in those pedestrian routes located in areas with enough population density over well-equipped central areas, so that the number of potential users justifies their investment and maintenance costs.

A comparative analysis of the vestibular apparatus in Epipliopithecus vindobonensis: Phylogenetic implications

Pliopithecoids are an extinct group of catarrhine primates from the Miocene of Eurasia. More than 50 years ago, they were linked to hylobatids due to some morphological similarities, but most subsequent studies have supported a stem catarrhine status, due to the retention of multiple plesiomorphic features (e.g., the ectotympanic morphology) relative to crown catarrhines. More recently, some morphological similarities to hominoids have been noted, raising the question of whether they could be stem members of this clade. To re-evaluate these competing hypotheses, we examine the morphology of the semicircular canals of the bony labyrinth of the middle Miocene pliopithecid Epipliopithecus vindobonensis. The semicircular canals are suitable to test between these hypotheses because (1) they have been shown to embed strong phylogenetic signal and reliably discriminate among major clades; (2) several potential hominoid synapomorphies have been identified previously in the semicircular canals; and (3) semicircular canal morphology has not been previously described for any pliopithecoid. We use a deformation-based (landmark-free) three-dimensional geometric morphometric approach to compare Epipliopithecus with a broad primate sample of extant and extinct anthropoids. We quantify similarities in semicircular canal morphology using multivariate analyses, reconstruct ancestral morphotypes by means of a phylomorphospace approach, and identify catarrhine and hominoid synapomorphies based on discrete characters. Epipliopithecus semicircular canal morphology most closely resembles that of platyrrhines and Aegyptopithecus due to the retention of multiple anthropoid symplesiomorphies. However, Epipliopithecus is most parsimoniously interpreted as a stem catarrhine more derived than Aegyptopithecus due to the possession of a crown catarrhine synapomorphy (i.e., the rounded anterior canal), combined with the lack of other catarrhine and any hominoid synapomorphies. Some similarities with hylobatids and atelids are interpreted as homoplasies likely related to positional behavior. The semicircular canal morphology of Epipliopithecus thus supports the common view that pliopithecoids are stem catarrhines.

Phylogeny of the Neotropical Pacman catfish genus Lophiosilurus (Siluriformes: Pseudopimelodidae)

Abstract Lophiosilurus is a monotypic genus represented by L. alexandri, a species endemic to the São Francisco river basin, Brazil. In previous phylogenetic analyses, the genus has been recovered as the sister group of Cephalosilurus. However, few species of Cephalosilurus or few characters were included in those studies. Thus, the current study aims to test the monophyletic hypothesis of the genera Lophiosilurus and Cephalosilurus with a more comprehensive phylogenetic analysis, including all Cephalosilurus species and a representative number of characters. Phylogenetic analyses of 18 terminal taxa (15 ingroups and three outgroups) were conducted based on a combined 75 character matrix, including 70 discrete morphological characters concerning osteology and neuroanatomy, four continuous characters, and the geometric morphometry of the head. The monophyly of the family Pseudopimelodidae was highly supported, and Cephalosilurus is synonymized with Lophiosilurus. The recovered phylogeny of the genus was (L. albomarginatus (L. nigricaudus (L. apurensis (L. fowleri, L. alexandri)))).

Synonymisation of the male-based ant genus

Although molecular data have proven indispensable in confidently resolving the phylogeny of many clades across the tree of life, these data may be inaccessible for certain taxa. The resolution of taxonomy in the ant subfamily Leptanillinae is made problematic by the absence of DNA sequence data for leptanilline taxa that are known only from male specimens, including the monotypic genus Phaulomyrma Wheeler & Wheeler. Focusing upon the considerable diversity of undescribed male leptanilline morphospecies, the phylogeny of 35 putative morphospecies sampled from across the Leptanillinae, plus an outgroup, is inferred from 11 nuclear loci and 41 discrete male morphological characters using a Bayesian total-evidence framework, with Phaulomyrma represented by morphological data only. Based upon the results of this analysis Phaulomyrma is synonymised with Leptanilla Emery, and male-based diagnoses for Leptanilla that are grounded in phylogeny are provided, under both broad and narrow circumscriptions of that genus. This demonstrates the potential utility of a total-evidence approach in inferring the phylogeny of rare extant taxa for which molecular data are unavailable and begins a long-overdue systematic revision of the Leptanillinae that is focused on male material.