Ancestral Character State Reconstruction Research Articles

Defining plant ecological specialists and generalists: Building a framework for identification and classification.

Specialization is a widespread but highly ambiguous and context-dependent ecological concept. This quality makes comparisons across related studies difficult and makes associated terms such as "specialist" and "generalist" scientifically obscure. Here, we present a metric-based framework to quantify specialization in 141 Quercus species using functional traits, biogeography, and species interactions. Rankings of specialization based on five metrics were used to answer questions about how specialization is used colloquially (i.e., individual species assessment by experts) and influenced by phylogenetics (Ancestral Character State Reconstruction, Automatic Shift Detection), biogeography (patterns of clustering by region and with climate), and species threat level (IUCN Red List). Metric-based ranking can be representative of specialization in a consistent and practical manner, correlating with IUCN Red List data, and the mean scores of individual expert assessments. Specialization is shown to be highly correlated with precipitation seasonality and only moderately influenced by evolutionary history. Data-deficient species were more likely to be highly specialized, and higher specialization was positively correlated with greater IUCN threat level. Frameworks for characterizing specialization and generalization can be done using metric ranking and can turn concepts that are often unclear into a definitive system. Metric-based rankings of specialization can also be used to reveal interesting insights about a clade's evolutionary history and geographic distribution when paired with the related phylogenetic and geographic data. Metric-based rankings can be applied to other systems and be a valuable tool for identifying species at risk and in need of conservation.

Trichomes’ Micromorphology and Their Evolution in Selected Species of Causonis (Vitaceae)

The Vitaceae genus Causonis is found in tropical to temperate climates from Asia to Australia, including the Pacific Islands. Rafinesque established the genus in 1930; however, Süssenguth classified it under Cayratia as a sect. Discypharia in 1953. The genus was resurrected in 2013 using morphological and genetic evidence. We herein provided insight into the diversity of trichomes’ micromorphology of selected species of this recently reinstated genus for taxonomical implication. Simple trichomes, representing non-glandular and unbranched trichomes, are only found in Causonis. Trichomes vary from straight, curved, hooked, appressed, pilose, to villous in different parts like branchlets, abaxial leaf surface, and adaxial leaf surface in different species. They also vary in the same plant from the young stage to the mature stage. Most species are pubescent when young, but a few species become nearly glabrous when they are mature. Significant variations can be observed in trichomes’ length between the species. Principal component analysis (PCA), based on the micromorphological traits, was carried out for the species delimitation. In Mesquite, ancestral character state reconstruction was used to examine evolutionary trends for trichomes on three different surfaces. The glabrous to sparsely pubescent state found on the branchlets and both leaf surfaces were found to be the ancestral state and, on the branchlets and both leaf surfaces, the villous hairs state was the derived state in the genus. The identification of Causonis species is greatly aided by trichomes morphology. Therefore, similar studies should be conducted on other Vitaceae genera to reveal the variety of trichomes found in the family.

Evolution and Biogeographic History of Rubyspot Damselflies (Hetaerininae: Calopterygidae: Odonata)

The damselflies Hetaerininae, a subfamily of Calopterygidae, comprise four genera distributed from North to South America: Hetaerina, Mnesarete, Ormenophlebia and Bryoplathanon. While several studies have focused on the intriguing behavioral and morphological modifications within Hetaerina, little of the evolutionary history of the group is well understood. Understanding the biogeographical history of Hetaerininae is further complicated by uncertainty in important geological events, such as the closure of the Central American Seaway (CAS). We generated a phylogenetic hypothesis to test the relationships and divergence times within Hetaerininae using IQtree and BEAST2 and found that Mnesarete and Ormenophlebia render Hetaerina paraphyletic. Reclassification of the genera within Hetaerininae is necessary based on our results. We also tested the fit to our dataset of two different hypotheses for the closure of CAS. Our results supported a gradual closure, starting in the Oligocene and ending in the Pliocene. Using Ancestral Character State Reconstruction, we found that the rubyspot, which is associated with higher fecundity in several species, was ancestral for Hetaerininae and subsequently lost four times. Estimates of diversification in association with the rubyspot are needed to understand the plasticity of this important character. Forest habitat was the ancestral state for Hetaerininae, with transitions to generalist species of Hetaerina found primarily in the Mesoamerican region. These results add to our understanding of the relationship between morphology, biogeography and habitat in a charismatic group of damselflies.

Re‐terrestrialization in the phylogeny of epiphytic plant lineages: Microsoroid ferns as a case study

AbstractThe fern family Polypodiaceae, with over 1600 species, is not only one of the most species‐rich families of ferns, but also a major contributor to the vascular epiphytic diversity throughout the tropics. Although the vast majority of species belonging to this family prefer to grow as epiphytes, several species colonize successfully rheophytic, lithophytic, and even terrestrial habitats. Here, we explore the hypothesis that non‐epiphytic habitat preferences, including terrestrial growth, evolved secondarily with epiphytes being the plesiomorphic habitat preference. The results of phylogenetic analyses, based on dense taxon sampling and four chloroplast DNA regions, were integrated with divergence time estimates and ancestral character state reconstructions to test these predictions. Both fossils and secondary calibration data were incorporated to obtain divergence time estimations. The results support the prediction of multiple transitions from epiphytic/lithophytic to terrestrial/rheophytic habitats occurring mainly in the Microsoreae lineage. The change in niche preferences coincides with niche colonization opportunities created by climatic fluctuations and geographical changes during the Oligocene and Miocene periods.

Climate drives anuran breeding phenology in a continental perspective as revealed by citizen‐collected data

AbstractAimAs anuran reproduction is generally linked to the availability of water, frogs and toads are particularly sensitive to climate. We tested the effect of climate on anuran reproductive phenology and daily activity by analysing temporal patterns of reproductive behaviour based on citizen‐collected observations.LocationBrazil.MethodsWe obtained vocalizations and photographs of frogs with inflated air sacs, as well as images of amplectant couples, nests, eggs or tadpoles in initial stages of development from iNaturalist. We analysed hourly patterns, seasonality and duration of the reproductive period using circular statistics in different climate types and tested phylogenetic signals. We analysed data for Bufonidae, Hylidae, Leptodactylidae and Phyllomedusidae in detail. We also reviewed relevant literature.ResultsAmong the 8478 (acoustic and photographic) records, 738 (8.7%) had evidence of reproduction with 284 acoustic records and 454 photographs, representing 184 taxa identified at the species level belonging to 16 families. Climate affected the period and duration of the reproductive season, as well as daily patterns of vocalization. These results were considered phylogenetically independent, as the reconstructions of ancestral character states did not suggest strong phylogenetic signals for temporal patterns of vocalization or reproduction. In Brazil, most frogs reproduce between October and January. Patterns were similar to the results of the literature review, however in the literature data, Bufonidae start reproduction 2 months earlier, and many Hylidae species reproduce until February. In general, frogs from warmer and drier climate regions had shorter and aggregated reproductive seasons, while in tropical monsoon climate they had a uniform temporal pattern.Main conclusionsAt the continental scale, reproductive phenology of anurans and their daily activity is affected by climatic conditions regardless of phylogeny. We found that community science can provide valuable information in Brazil that can be harnessed to monitor effects of climate change on amphibian reproduction.

Incomplete lineage sorting and reticulate evolution mask species relationships in Brunelliaceae, an Andean family with rapid, recent diversification.

To date, phylogenetic relationships within the monogeneric Brunelliaceae have been based on morphological evidence, which does not provide sufficient phylogenetic resolution. Here we use target-enriched nuclear data to improve our understanding of phylogenetic relationships in the family. We used the Angiosperms353 toolkit for targeted recovery of exonic regions and supercontigs (exons + introns) from low copy nuclear genes from 53 of 70 species in Brunellia, and several outgroup taxa. We removed loci that indicated biased inference of relationships and applied concatenated and coalescent methods to infer Brunellia phylogeny. We identified conflicts among gene trees that may reflect hybridization or incomplete lineage sorting events and assessed their impact on phylogenetic inference. Finally, we performed ancestral-state reconstructions of morphological traits and assessed the homology of character states used to define sections and subsections in Brunellia. Brunellia comprises two major clades and several subclades. Most of these clades/subclades do not correspond to previous infrageneric taxa. There is high topological incongruence among the subclades across analyses. Phylogenetic reconstructions point to rapid species diversification in Brunelliaceae, reflected in very short branches between successive species splits. The removal of putatively biased loci slightly improves phylogenetic support for individual clades. Reticulate evolution due to hybridization and/or incomplete lineage sorting likely both contribute to gene-tree discordance. Morphological characters used to define taxa in current classification schemes are homoplastic in the ancestral character-state reconstructions. While target enrichment data allows us to broaden our understanding of diversification in Brunellia, the relationships among subclades remain incompletely understood.

Phylogenomic loci define the generic boundaries of Gochnatieae and improve resolution at the species level in Moquiniastrum (Compositae)

Understanding the evolution of the tribe Gochnatieae (Compositae) has been the subject of considerable effort in the past decade. This is due to the key position of this tribe in the phylogeny of the sunflower family and the corresponding implications for biogeographic and morphological evolution of Compositae. Previous studies have confirmed the monophyly of this tribe as well as most of the genera that belong to it. However, phylogenetic resolution of Gochnatieae at both the genus- and species-level has remained poor. A subset of new phylogenomic loci used in this study has proven effective and has improved phylogenetic resolution in this group. The results of this work demonstrate Gochnatieae is a well-supported clade comprised of nine genera (Anastraphia, Cnicothamnus, Cyclolepis, Gochnatia, Moquiniastrum, Nahuatlea, Pentaphorus, Richterago, Tehuasca). One recently described genus, Vickia, was not included in this study; but its placement in Gochnatieae as a tenth genus in the tribe is well-justified. The monospecific Cyclolepis, which had been circumscribed within the tribe since its inception but was subsequently removed and designated as incertae sedis since 2014, is also shown to belong to Gochnatieae. We confirmed the monophyletic Moquiniastrum with two well-supported subclades. Ancestral area reconstruction analyses show that Gochnatieae originated in Eastern South America about 53 my. Apparently, except for Cyclolepis and Richterago, the ancestors of the other genera of Gochnatieae originated about 44 my from an area that now corresponds to the central Andes. The presence of the genera in the Chaco phytogeographic province, central Chile, and Mexico-United States-Caribbean is a result of dispersal from the central Andes. The ancestral distribution of Moquiniastrum corresponds to a large area comprising Eastern South America and the current central Andes, about 32 my. Ancestral character state reconstruction that included four characters indicates several states associated with complex plant reproductive biology such as gynodioecy, gynomonoecy, and polygamodioecy are derived in Gochnatieae as are heterogamous capitula (in Moquiniastrum and Richterago), dimorphic and subdimorphic corollas (in Cnicothamnus, Moquiniastrum, and Richterago), and the presence of marginal female corollas (in Moquiniastrum and Richterago). Within Moquiniastrum, two subclades (Densicephalum and Polymorphum) exhibit divergent patterns of trait evolution associated with these reproductive characters which suggests this genus can serve as a model to understand the sexual system evolution in plants.

Hydathodes in ferns: their phylogenetic distribution, structure and function.

Ferns are the second largest group of vascular plants and are distributed nearly worldwide. Although ferns have been integrated into some comparative ecological studies focusing on hydathodes, there is a considerable gap in our understanding of the functional anatomy of these secretory tissues that are found on the vein endings of many fern leaves. In this study, we aimed to investigate the phylogenetic distribution, structure and function of fern hydathodes. We performed a global review on fern hydathodes and their phylogenetic distribution, carried out an ancestral character state reconstruction, and studied the structure, guttation and elemental composition of salt residues of eight species, and the diurnal patterns of xylem pressure of two species. Hydathodes are known from 1189 fern species, 92 genera and 19 families of 2 orders, Equisetales and Polypodiales. Stochastic character mapping indicated multiple gains and losses of hydathodes at the genus level, occurring especially during the last 50 million years of fern evolution. Hydathodes were located on the adaxial leaf surface and characterized by a cytoplasm-rich, pore-free epidermis, and became functional for several weeks after nearly complete leaf expansion. In two species, positive xylem pressure built up at night, potentially facilitating guttation. Guttation fluid was rich in Ca and often Si, but also contained P, Mg, Na and Al. Stochastic character mapping and the structural and functional diversity of hydathodes indicate multiple origins, and their presence/absence in closely related taxa implies secondary losses during fern evolution. Positive xylem pressure and high air humidity play an important role as drivers of guttation. Hydathodes may contribute to the regulation of leaf nutrient stoichiometry by the release of excessive compounds and minerals other than waste products, but the presence of essential chemical elements in salt residues also indicates possible leakage.

Mitogenomic phylogeny of Typhlocybinae (Hemiptera: Cicadellidae) reveals homoplasy in tribal diagnostic morphological traits.

The subfamily Typhlocybinae is a ubiquitous, highly diverse group of mostly tiny, delicate leafhoppers. The tribal classification has long been controversial and phylogenetic methods have only recently begun to test the phylogenetic status and relationships of tribes. To shed light on the evolution of Typhlocybinae, we performed phylogenetic analyses based on 28 newly sequenced and 19 previously sequenced mitochondrial genomes representing all currently recognized tribes. The results support the monophyly of the subfamily and its sister‐group relationship to Mileewinae. The tribe Zyginellini is polyphyletic with some included genera derived independently within Typhlocybini. Ancestral character state reconstruction suggests that some morphological characters traditionally considered important for diagnosing tribes (presence/absence of ocelli, development of hind wing submarginal vein) are homoplastic. Divergence time estimates indicate that the subfamily arose during the Middle Cretaceous and that the extant tribes arose during the Late Cretaceous. Phylogenetic results support establishment of a new genus, Subtilissimia Yan & Yang gen. nov., with two new species, Subtilissimia fulva Yan & Yang sp. nov. and Subtilissimia pellicula Yan & Yang sp. nov.; but indicate that two previously recognized species of Farynala distinguished only by the direction of curvature of the processes of the aedeagus are synonyms, that is, Farynala dextra Yan & Yang, 2017 equals Farynala sinistra Yan & Yang, 2017 syn. nov. A key to tribes of Typhlocybinae is provided.

Ecological speciation by sympatric host shifts in a clade of herbivorous sea slugs, with introgression and localized mitochondrial capture between species

Host shifting in insect-plant systems was historically important to the development of ecological speciation theory, yet surprisingly few studies have examined whether host shifting drives diversification of marine herbivores. When small-bodied consumers feed and also mate on a preferred host, disruptive selection can split a population into host races despite gene flow. Support for host shifts is notably lacking for invertebrates associated with macroalgae, where the scale of dispersal by planktonic larvae often far exceeds the grain of host patchiness, and adults are typically less specialized than terrestrial herbivores. Here, we present a candidate example of ecological speciation in a clade of sea slugs that primarily consume green algae in the genus Caulerpa, including highly invasive species. Ancestral character state reconstructions supported ‘sea grapes’ (C. racemosa, C. lentillifera) as the ancestral host for a tropical radiation of 12 Elysia spp., with one shift onto alternative Caulerpa spp. in the Indo-Pacific. A Caribbean radiation of three species included symaptric host shifts to Rhipocephalus brevicaulis in the ancestor of E. pratensis Ortea & Espinosa, 1996, and to C. prolifera in E. hamanni Krug, Vendetti & Valdes 2016, plus a niche expansion to a range of Caulerpa spp. in E. subornata Verrill, 1901. All three species are broadly sympatric across the Caribbean but are host-partitioned at a fine grain, and distinct by morphology and at nuclear loci. However, non-recombining mtDNA revealed a history of gene flow between E. pratensis and E. subornata: COI haplotypes from E. subornata were 10.4% divergent from E. pratensis haplotypes from four sites, but closely related to all E. pratensis haplotypes sampled from six Bahamian islands, indicating historical introgression and localized “mitochondrial capture.” Disruptive selective likely fueled divergence and adaptation to distinct host environments, indicating ecological speciation may be an under-appreciated driver of diversification for marine herbivores as well as epibionts and other resource specialists.

Comparative genomic analysis of Thermus provides insights into the evolutionary history of an incomplete denitrification pathway.

Biological denitrification is a crucial process in the nitrogen biogeochemical cycle, and Thermus has been reported to be a significant heterotrophic denitrifier in terrestrial geothermal environments. However, neither the denitrification potential nor the evolutionary history of denitrification genes in the genus Thermus or phylum Deinococcota is well understood. Here, we performed a comparative analysis of 23 Thermus genomes and identified denitrification genes in 15 Thermusstrains. We confirmed that Thermus harbors an incomplete denitrification pathway asnone of the strains contain the nosZ gene. Ancestral character state reconstructions and phylogenetic analyses showed that narG, nirS, and norB genes were acquired by the last common ancestor of Thermales and were inherited vertically. In contrast, nirK of Thermales was acquired via two distinct horizontal gene transfers from Proteobacteria to the genus Caldithermus and from an unknown donor to the common ancestor of all known Thermus species except Thermus filiformis. This study expands our understanding of the genomic potential for incomplete denitrification in Thermus, revealinga largely vertical evolutionary history of the denitrification pathway in the Thermaceae, and supportingthe important role for Thermus as an important heterotrophic denitrifier in geothermal environments.

The Evolution of Cytogenetic Traits in Cuscuta (Convolvulaceae), the Genus With the Most Diverse Chromosomes in Angiosperms.

Karyotypes are characterized by traits such as chromosome number, which can change through whole-genome duplication and dysploidy. In the parasitic plant genus Cuscuta (Convolvulaceae), chromosome numbers vary more than 18-fold. In addition, species of this group show the highest diversity in terms of genome size among angiosperms, as well as a wide variation in the number and distribution of 5S and 35S ribosomal DNA (rDNA) sites. To understand its karyotypic evolution, ancestral character state reconstructions were performed for chromosome number, genome size, and position of 5S and 35S rDNA sites. Previous cytogenetic data were reviewed and complemented with original chromosome counts, genome size estimates, and rDNA distribution assessed via fluorescence in situ hybridization (FISH), for two, seven, and 10 species, respectively. Starting from an ancestral chromosome number of x = 15, duplications were inferred as the prevalent evolutionary process. However, in holocentric clade (subgenus Cuscuta), dysploidy was identified as the main evolutionary mechanism, typical of holocentric karyotypes. The ancestral genome size of Cuscuta was inferred as approximately 1C = 12 Gbp, with an average genome size of 1C = 2.8 Gbp. This indicates an expansion of the genome size relative to other Convolvulaceae, which may be linked to the parasitic lifestyle of Cuscuta. Finally, the position of rDNA sites varied mostly in species with multiple sites in the same karyotype. This feature may be related to the amplification of rDNA sites in association to other repeats present in the heterochromatin. The data suggest that different mechanisms acted in different subgenera, generating the exceptional diversity of karyotypes in Cuscuta.

Evolution of the Anther Gland in Early-Branching Papilionoids (ADA Clade, Papilionoideae, Leguminosae).

Papilionoideae is the most diverse subfamily of Leguminosae, especially in terms of floral morphology. The ADA clade shows some exciting floral features among papilionoids, such as anther glands. However, the evolution of the anther glands in such early-branching papilionoids remains unknown. Thus, we compared the occurrence, distribution, morphology, and evolutionary history of the anther glands in species of the ADA clade. Floral buds and/or flowers in 50 species were collected from herbarium specimens and investigated using scanning electron and light microscopy and reconstruction of ancestral character states. The anther apex has a secretory cavity, secretory duct, and phenolic idioblast. The lumen shape of the cavity and duct is closely related to the shape of the anther apex. The oval lumen is located between two thecae, the spherical lumen in the prominent anther apex and the elongated lumen in anthers with a long apex. The occurrence of cavities/ducts in the anther in only two phylogenetically closely related subclades is a unifying character -state. The floral architecture is not correlated with cavity/ducts in the anther but is possibly related to the type of pollinator. Future research needs to combine floral morphology and pollination systems to understand the evolution of floral designs and their diversification.

Molecular and Morphological Analysis Supports the Separation ofRobrichiaas a Genus Distinct fromEnterolobium(Leguminosae: Caesalpinioideae: Mimosoid Clade)

Abstract—As currently circumscribed, the legume genusEnterolobiumcomprises 11 species in two sections:E.sect.EnterolobiumandE.sect.Robrichia, with an overall distribution from Mexico to Argentina and a centre of diversity in Amazonia. In the absence of the characteristic indehiscent fruits,Enterolobiumis difficult to distinguish from other genera in the ingoid clade, includingAlbiziasensu lato,Samanea, andCathormion. Previous phylogenetic studies which have includedEnterolobiumhave sampled few species of the genus, leaving questions about its monophyly and interspecific relationships. Here we evaluate the circumscription ofEnterolobiumand its two infrageneric taxa, their phylogenetic placement in the ingoid clade, and interspecific relationships within the two sections of the genus. Our study includes allEnterolobiumspecies, and analyses of nuclear (ITS and ETS) and plastid (trnD-TandtrnL-F) molecular regions, and morphology. Bayesian inference, maximum parsimony, and maximum likelihood analyses show thatEnterolobium, as presently circumscribed, is not monophyletic. The genus is divided into two well-supported independent clades, corresponding to the two previously recognized sections; one of them is sister to the genusLeucochloronand is here raised to generic rank as the genusRobrichia, with three species,Robrichia glaziovii,Robrichia oldemaniiandRobrichia schomburgkii. The genusEnterolobiumconsequently now comprises eight species and is more closely related to a clade composed ofAlbiziaandFalcataria. This new arrangement is reinforced by morphological synapomorphies recovered by ancestral character state reconstructions. Indumentum type, the number of pinnae pairs per bipinnate leaf and leaflet pairs per pinna, inflorescence type, and fruit shape characterizeRobrichia, while the recognition ofEnterolobiumremains based on fruit traits. We also provide the formal lectotypifications forE. contortisiliquum,E. cyclocarpum,E. gummiferum,E. maximum,E. monjollo, andR. schomburgkii.

Phylogenetics of Serjania (Sapindaceae-Paullinieae), with emphasis on fruit evolution and the description of a new species from Michoacán, Mexico

Serjania is among the largest Neotropical genera of Sapindaceae and comprises ∼240 species. Traditionally, the genus has been defined by its schizocarpic fruits separating into three distally winged mericarps. However, recent phylogenetic studies have revealed that fruit type is not consistent within the genus and that it also includes a few species having capsular fruits that were previously placed in other genera. A phylogenetic analysis is presented based on nuclear ITS and chloroplast trnL-F sequences. The ingroup consisted of 48 species broadly sampled from across the geographic range and taxonomic spectrum of Serjania and focusing on species with atypical fruits. An ancestral character state reconstruction of fruit type was performed and demonstrated that the ancestral fruit type of the genus is strongly supported to be a schizocarp, but there are at least five independent derivations to capsular fruits and at least one transition back from capsules to schizocarps. Also, transitions from winged to wingless mericarps have occurred at least twice. Infrageneric classification is problematic and all of the sections that were represented by more than one species come out as poly- or paraphyletic. Our limited sample of taxa precludes the possibility of a new classification at present and a broader phylogenetic sample of the genus will be needed to understand relationships and determine which lineages warrant formal recognition. In addition to the phylogenetic study, Serjania frutescens, a new capsular-fruited species from Michoacán, Mexico, is described, illustrated, and compared with its putative closest relatives. A key is provided to differentiate this species from other Mexican Serjania with capsular fruits, and new synonymies and lectotypifications associated with capsular-fruited Serjania tortuosa and S. sonorensis are established. Phylogenetic results suggest that these two are cryptic species and at least superficially indistinguishable by morphology.

Himalayan orogeny and monsoon intensification explain species diversification in an endemic ginger (Hedychium: Zingiberaceae) from the Indo-Malayan Realm

The Indo-Malayan Realm is a biogeographic realm that extends from the Indian Subcontinent to the islands of Southeast Asia (Malay Archipelago). Despite being megadiverse, evolutionary hypotheses explaining taxonomic diversity in this region have been rare. Here, we investigate the role of geoclimatic events such as Himalayan orogeny and monsoon intensification in the diversification of the ginger-lilies (Hedychium J.Koenig: Zingiberaceae). We first built a comprehensive, time-calibrated phylogeny of Hedychium with 75% taxonomic and geographic sampling. We found that Hedychium is a very young lineage that originated in Northern Indo-Burma, in the Late Miocene (c. 10.6 Ma). This was followed by a late Neogene and early Quaternary diversification, with multiple dispersal events to Southern Indo-Burma, Himalayas, Peninsular India, and the Malay Archipelago. The most speciose clade IV i.e., the predominantly Indo-Burmese clade also showed a higher diversification rate, suggesting its recent rapid radiation. Our divergence dating and GeoHiSSE results demonstrate that the diversification of Hedychium was shaped by both the intensifications in the Himalayan uplift as well as the Asian monsoon. Ancestral character-state reconstructions identified the occurrence of vegetative dormancy in both clades I and II, whereas the strictly epiphytic growth behavior, island dwarfism, lack of dormancy, and a distinct environmental niche were observed only in the predominantly island clade i.e., clade III. Finally, we show that the occurrence of epiphytism in clade III corresponds with submergence due to sea-level changes, suggesting it to be an adaptive trait. Our study highlights the role of recent geoclimatic events and environmental factors in the diversification of plants within the Indo-Malayan Realm and the need for collaborative work to understand biogeographic patterns within this understudied region. This study opens new perspectives for future biogeographic studies in this region and provides a framework to explain the taxonomic hyperdiversity of the Indo-Malayan Realm.

Dense sampling of taxa and characters improves phylogenetic resolution among deltocephaline leafhoppers (Hemiptera: Cicadellidae: Deltocephalinae)

AbstractWe analysed a phylogenomic dataset comprising 730 terminal taxa and >160,000 nucleotide positions obtained using anchored hybrid enrichment of genomic DNA for a sample of deltocephaline leafhoppers and outgroups. Maximum likelihood analyses of concatenated nucleotide and amino acid sequences as well as coalescent gene tree analysis, yielded well‐resolved phylogenetic estimates that were highly congruent with most branches receiving maximum support. Some topological incongruence occurs among the trees resulting from different analyses, mainly distributed among very short branches at intermediate levels in the phylogeny, pertaining to relationships among some tribes and multi‐tribe lineages restricted to particular continents. Coalescent gene tree analysis revealed extensive gene tree conflict at these nodes, suggesting that certain relationships may remain difficult to resolve consistently even with genome‐scale datasets and dense taxon sampling. Ancestral character state reconstruction of feeding preference indicates that grass specialization has been acquired three or fewer times in some highly diverse but relatively derived lineages. Molecular divergence time estimation suggests that the earliest divergences in the subfamily occurred during the Cretaceous but that most modern tribes did not appear until after the Cretaceous–Palaeogene boundary and that grass specialist lineages generally diversified and spread at the same time as grasslands were becoming widespread globally. These analyses also show a high level of global biogeographic structure, with several large lineages of primarily arboreal deltocephalines restricted to particular regions or continents and long‐distance dispersal among continents occurring primarily among grass‐specialist lineages. The results represent a major improvement over previous analyses of this diverse subfamily, which were based on 152 or fewer taxa and data from morphology and partial sequences of two genes.

Phylogeny and secondary sexual trait evolution in Schizocosa wolf spiders (Araneae, Lycosidae) shows evidence for multiple gains and losses of ornamentation and species delimitation uncertainty

Members of the Nearctic spider genus Schizocosa Chamberlin, 1904 have garnered much attention in behavioral studies and over many decades, a number of species have developed as model systems for investigating patterns of sexual selection and multimodal communication. Many of these studies have employed a comparative approach using putative, but not rigorously tested, sister species pairs that have distinctive morphological traits and attendant behaviors. Despite past emphasis on the efficacy of these presumably comparative-based studies of closely related species, generating a robust phylogenetic hypothesis for Schizocosa has been an ongoing challenge. Here, we apply a phylogenomic approach using anchored hybrid enrichment to generate a data set comprising over 400 loci representing a comprehensive taxonomic sample of 23 Nearctic Schizocosa. Our sampling also includes numerous outgroup lycosid genera that allow for a robust evaluation of genus monophyly. Based on analyses using concatenation and coalescent-based methods, we recover a well-supported phylogeny that infers the following: 1) The New World Schizocosa do not form a monophyletic group; 2) Previous hypotheses of North American species require reconsideration along with the composition of species groups; 3) Multiple longstanding model species are not genealogically exclusive and thus are not “good” species; 4) This updated phylogenetic framework establishes a new working paradigm for studying the evolution of characters associated with reproductive communication and mating. Ancestral character state reconstructions show a complex pattern of homoplasy that has likely obfuscated previous attempts to reconstruct relationships and delimit species. Important characters presumably related to sexual selection, such as foreleg pigmentation and dense bristle formation, have undergone repeated gain and loss events, many of which have led to increased morphological divergence between sister-species. Evaluation of these traits in a comparative framework illuminates how sexual selection and natural selection influence character evolution and provides a model for future studies of multimodal communication evolution and function.

Taxonomy, phylogeny, molecular dating and ancestral state reconstruction of Xylariomycetidae (Sordariomycetes)

Xylariomycetidae (Ascomycota) is a highly diversified group with variable stromatic characters. Our research focused on inconspicuous stromatic xylarialean taxa from China, Italy, Russia, Thailand and the United Kingdom. Detailed morphological descriptions, illustrations and combined ITS-LSU-rpb2-tub2-tef1 phylogenies revealed 39 taxa from our collections belonging to Amphisphaeriales and Xylariales. A new family (Appendicosporaceae), five new genera (Magnostiolata, Melanostictus, Neoamphisphaeria, Nigropunctata and Paravamsapriya), 27 new species (Acrocordiella photiniicola, Allocryptovalsa sichuanensis, Amphisphaeria parvispora, Anthostomella lamiacearum, Apiospora guiyangensis, A. sichuanensis, Biscogniauxia magna, Eutypa camelliae, Helicogermslita clypeata, Hypocopra zeae, Magnostiolata mucida, Melanostictus longiostiolatus, M. thailandicus, Nemania longipedicellata, N. delonicis, N. paraphysata, N. thailandensis, Neoamphisphaeria hyalinospora, Neoanthostomella bambusicola, Nigropunctata bambusicola, N. nigrocircularis, N. thailandica, Occultitheca rosae, Paravamsapriya ostiolata, Peroneutypa leucaenae, Seiridium italicum and Vamsapriya mucosa) and seven new host/geographical records are introduced and reported. Divergence time estimates indicate that Delonicicolales diverged from Amphisphaeriales + Xylariales at 161 (123–197) MYA. Amphisphaeriales and Xylariales diverged 154 (117–190) MYA with a crown age of 127 (92–165) MYA and 147 (111–184) MYA, respectively. Appendicosporaceae (Amphisphaeriales) has a stem age of 89 (65–117) MYA. Ancestral character state reconstruction indicates that astromatic, clypeate ascomata with aseptate, hyaline ascospores that lack germ slits may probably be ancestral Xylariomycetidae having plant-fungal endophytic associations. The Amphisphaeriales remained mostly astromatic with common septate, hyaline ascospores. Stromatic variations may have developed mostly during the Cretaceous period. Brown ascospores are common in Xylariales, but they first appeared in Amphisphaeriaceae, Melogrammataceae and Sporocadaceae during the early Cretaceous. The ascospore germ slits appeared only in Xylariales during the Cretaceous after the divergence of Lopadostomataceae. Hyaline, filiform and apiospores may have appeared as separate lineages, providing the basis for Xylariaceae, which may have diverged independently. The future classification of polyphyletic xylarialean taxa will not be based on stromatic variations, but the type of ring, the colour of the ascospores, and the presence or absence or the type of germ slit.

Morphological characters in light of new molecular phylogenies: the caudal-fin skeleton of Ovalentaria.

The Ovalentaria is a taxon of teleosts that has been proposed based on molecular analyses only. Previously widely separated families are assembled in this taxon. For the first time, the Ovalentaria are analysed using a comparative morphological approach. The caudal-fin skeleton of 355 species covering all 48 ovalentarian families are examined in cleared and stained specimens, µCT datasets and X-ray images as well as from the literature. A total of 38 morphological characters are evaluated and used for ancestral character state reconstructions and phylogenetic analyses. Results provide hypotheses for a scenario of the evolution of the caudal-fin skeleton and its ground plan in Ovalentaria. An evolutionary trend towards the reduction of skeletal elements in the caudal fin is observed. Connections between the evolution of the caudal-fin skeleton and modes of locomotion found in ovalentarian taxa are discussed. Phylogenetic analyses based on the caudal-fin morphology provide topologies for intra-ovalentarian relationships that largely agree with molecular hypotheses.