Ancestral Character State Research Articles

Linguistic homoplasy and phylogeny reconstruction. The cases of Lezgian and Tsezic languages (North Caucasus)

Abstract This paper deals with the problem of linguistic homoplasy (parallel or backward development), how it can be detected, what kinds of linguistic homoplasy can be distinguished and which varieties of the phenomenon are the most deleterious for the reconstruction of language phylogeny. It is proposed that language phylogeny reconstruction should consist of two main stages. Firstly, a strict consensus tree should be built on the basis of high-quality input data elaborated with the help of the main phylogenetic methods (such as Neighbor-joining, Bayesian MCMC, and Maximum parsimony), and ancestral character states, allowing us to reveal a certain number of homoplastic characters. Secondly, after the detected instances of homoplasy are eliminated from the input matrix, the consensus tree is to be compiled again. It is expected that after homoplastic optimization it will be possible to better resolve individual “problem clades”, and generally the homoplasy-optimized phylogeny should be more robust than the tree constructed initially. The proposed procedure is tested on the 110-item Swadesh wordlists of the Lezgian and Tsezic groups. The Lezgian and Tsezic results generally support theoretical expectations. The MLN (minimal lateral network) method, currently implemented in the LingPy software, is a helpful tool for the detection of linguistic homoplasy.

Congruence between comparative morphology and molecular phylogenies: the final stage of evolution of the skeletal characters of male genitalia in the subtribe Polyommatina (Lepidoptera, Lycaenidae). Part 2. Uncus and subunci

The morphology of the male terminalia (uncus and subunci) was studied in 39 species of the subtribe Polyommatina (Lycaenidae). Traces of nine established characters were mapped on the molecular phylogeny of the subtribe proposed by Talavera et al. (2013). The ancestral character states for these morphological structures were established in Plebejus s. l. and Polyommatus s. l. According to the time calibration of the proposed molecular tree, the studied skeletal structures (uncus and subunci) evolved very slowly within the last million years. It was shown that the similar shape of the uncus and subuncus branches arose independently and underwent parallel morphological transformations. In particular, wide and, as a rule, ventrally directed uncus lobes arose independently in Polyommatus s. str., Alpherakya, Glabroculus, and Aricia. Yet, in various evolutionary lines of Polyommatina similar changes in the length ratio of the proximal and distal parts of the subunci as well as in the shape of their apical hooks were observed. The character states marking the groups Polyommatus s. str. and Aricia were established. The evolutionary changes in the uncus morphology were highly non-uniform in the various monophyletic lines of Polyommatina. As a result, representatives of some “old” lines have more specialized features of the uncus morphology in comparison with those belonging to “young” lines that retain ancestral character states. In Polyommatina, the transformation of the male terminalia was directed toward development of the most effective mechanism of female fixation during copulation. This goal was reached in similar ways in various evolutionary lines of Polyommatina, which explains numerous parallelisms in morphology of these structures.

Tandem duplication of two tRNA genes in the mitochondrial genome of Tagiades vajuna (Lepidoptera: Hesperiidae)

To explore the debated phylogenetic relationship of two Hesperiidae subfamilies, Pyrginae and Eudaminae, and contribute to the understanding of the evolution of mitogenomic architecture in butterflies, we sequenced the complete mitogenome of Tagiades vajuna. The mitogenome is a typical circular duplex molecule of 15,359 bp. Apart from the standard 22 tRNAs, it has a tandem duplication of trnS(AGN) and trnE, which is unique in lepidopteran insects. Comparison with Ctenoptilum vasava indicates that the trnS1 duplication is not an ancestral state shared with other species of Tagiadini. Independent origin of the trnS1 duplications was further confirmed by the reconstruction of the ancestral character state based on the topology of the phylogram. Furthermore, comparative analysis of mitogenomes with and without tRNA duplications indicates that tRNA duplication does not alter the codon usage pattern. The mitogenome has negative AT- and GC-skews, and it is highly A+T-biased (79.7%). The AT-rich (or control) region (283 bp) contains ATAGA and ATTTA motifs. Regarding the phylogenetic analysis, we found that removal of the third codon position (3CP) from datasets used for the mitochondrial phylogenomics of Hesperiidae is likely to produce results that are more consistent: Pyrginae were rendered paraphyletic by Eudaminae in both analyses of the dataset from which the 3CP was removed (13 PCGs + all RNAs), but inclusion of the 3CP resulted in a destabilized topology, resulting in both monophyly and polyphyly. We conclude that even shallow-phylogenies of insects should pay close attention to compositional and mutational biases in mitogenomes.

Phylogeny, historical biogeography and characters evolution of the drought resistant fern Pyrrosia Mirbel (Polypodiaceae) inferred from plastid and nuclear markers

Pyrrosia s.l. comprises ca. 60 species with a disjunct Africa/Asia and Australia distribution. The infrageneric classification of Pyrrosia s.l. is controversial based on the phylogenetic analyses of chloroplast markers and morphology. Based on the expanded taxon sampling of Pyrrosia s.l. (51 species), we investigated its phylogeny, biogeography, character evolution and environmental adaptation by employing five chloroplastid markers (rbcL, matK, psbA-trnH, and rps4 + rps4-trnS) and one single (low)-copy nuclear gene, LEAFY. Pyrrosia s.l. was divided into six major clades and eight subclades. Reticulate evolution was revealed both among clades and among species in Pyrrosia s.l. Ancestral character state optimization revealed high levels of homoplastic evolution of the diagnostic characters in Pyrrosia s.l., while the crassulacean acid metabolism pathway seems to have an independent origin. Molecular dating and biogeographic diversification analyses suggested that Pyrrosia s.l. originated no later than the Oligocene and the main clades diversified during the Oligocene and Miocene, with southern Asia, the Indo-China Peninsula and southwestern and southern China as the most likely ancestral areas. Transoceanic long-distance dispersal, rather than vicariance, contributed to the intercontinental disjunction. Diversification scenarios of Pyrrosia s.l. under geological movements and climate fluctuation are also discussed.

Towards resolving the evolutionary history of Caucasian pears (Pyrus, Rosaceae)—Phylogenetic relationships, divergence times and leaf trait evolution

AbstractWith approximately 25 endemic species, the genus Pyrus (pears) is highly diverse in the Caucasus ecoregion. The majority of Caucasian pears inhabit xerophytic open woodlands or similar habitats, to which they display morphological adaptations, such as narrow leaves. The other species, both Caucasian and non‐Caucasian taxa, mainly inhabit mesophytic forests and display broad leaves. Using a representative taxon sampling of Pyrus from the Caucasus, Europe and Asia, we reconstruct phylogenetic relationships in the genus based on multiple plastid regions. We also estimate the divergence times of major clades in Pyrus, reconstruct the evolution of leaf shapes, and discuss the emergence of xeromorphic leaf traits. Our results confirm the monophyly of Pyrus and the existence of two major clades: (a) an E Asian clade with a crown group age of 15.7 (24.02–8.37 95% HPD) My, and (b) a W Eurasian clade that comprises species from Europe, SW Asia and the Caucasus and that displays a slightly younger crown group of 12.38 (19.02–6.41 95% HPD) My. The existing infrageneric classification of Pyrus was found partially incongruent with the inferred phylogenetic trees. Several currently accepted species were not recovered as monophyletic, indicating that current species limits require re‐evaluation. Ancestral character state reconstructions revealed several independent transitions from broad‐ to narrow‐shaped leaves in Pyrus, probably via intermediate‐shaped leaves.

Phylogenomics of Andropogoneae (Panicoideae: Poaceae) of Mainland Southeast Asia

The grass tribe Andropogoneae is distributed in warm regions around the globe but has been poorly studied in mainland Southeast Asia. This is particularly true for the cosmopolitan genera Andropogon and Schizachyrium, with several species that appear to be narrowly distributed in this region. Additionally, lesser-known species in the genera Hemisorghum, Kerriochloa, and Pseudosorghum also occur inmainland Southeast Asia. A phylogeny is needed to address questions of taxonomy and trait evolution. Whole chloroplast genomes of Andropogoneae species and two outgroup species of Garnotia (tribe Arundinelleae) were analyzed using maximum likelihood (ML) and Bayesian inference (BI). Ancestral character states were reconstructed usingML for four morphological characters key to Andropogon and Schizachyrium identification.Apreviously-unidentified clade of Southeast Asian endemic taxa is found, including one species formerly classified in Andropogon. Other Southeast Asian taxa fall in an unresolved grade outside the major radiation of the tribe. Andropogon and Schizachyrium are both polyphyletic. Convergent evolution and reversal of characters are common throughout Andropogoneae. Addition of species from mainland Southeast Asian finds unexpected phylogenetic diversity. Southeast Asian Schizachyrium sanguineum forms two separate clades, which could reflect cryptic species differentiation, hybridization, introgression, or some combination.

Phylogeny and evolution of achenial trichomes in theLucilia‐group (Asteraceae: Gnaphalieae) and their systematic significance

AbstractThe Gnaphalieae (Asteraceae) are a cosmopolitan tribe with around 185 genera and 2000 species. The New World is one of the centers of diversity of the tribe with 24 genera and over 100 species, most of which form a clade called theLucilia‐group with 21 genera. However, the generic classification of theLucilia‐group has been controversial with no agreement on delimitation or circumscription of genera. Especially controversial has been the taxonomic value of achenial trichomes and molecular studies have shown equivocal results so far. The major aims of this paper are to provide a nearly complete phylogeny of theLucilia‐group at generic level and to discuss the evolutionary trends and taxonomic significance of achenial trichome morphology. We conducted a phylogenetic analysis of the New World Gnaphalieae with nrDNA (ETS, ITS) sequence data from a sampling of 18 genera of theLucilia‐group and utilized these results to examine morphological evolution of achenial trichome types and presence of apical myxogenic cells. Seven well‐supported subclades can be recognized within theLucilia‐group (L1–L7). These results support Brazilian and AndeanBerroa,Facelis,Lucilia, andMicropsisforming a clade (L1), the inclusion of ChileanLuciliaunderBelloa(L2), the monophyly ofStuckertiella+Gamochaeta+Gamochaetopsis(L3),Chevreulia+Cuatrecasasiella(L4) andAntennaria(L5) excludingAntennaria linearifolia, which is resolved in a monophyletic group together withJalcophila,LoricariaandMniodes(L6), and the recognition ofGnaphaliothamnus(L7) removed from Brazilian taxa ofChionolaena(L2). Ancestral character state reconstruction of achenial trichome morphology suggests that clades are homogeneous in terms of trichome type, but with exceptions that make it highly homoplastic. Conversely, our results suggest that the presence of myxogenic apical cells is less homoplastic and that closely related species tend to resemble each other more than expected under random variation.

The evolutionary history of Senna ser. Aphyllae (Leguminosae–Caesalpinioideae), an endemic clade of southern South America

In the legume genus Senna, series Aphyllae includes seven species of leafless shrubs and subshrubs from arid, semiarid and xerophilous areas of three different biogeographic subregions in southern South America. In this study, we investigated the evolutionary history of Aphyllae in a molecular phylogenetic framework. We reconstructed phylogenetic relationships among Aphyllae species based on DNA sequence data of four plastid (rpS16, rpL16, matK, trnL-F) and one nuclear (ITS) region from 23 accessions, analyzed with parsimony, Bayesian and maximum likelihood methods. We inferred the evolutionary and biogeographic history estimating divergence times and reconstructing ancestral character states and ancestral areas of distribution. Series Aphyllae was found to be monophyletic, and the taxa formed two main clades: Clade A gathering S. aphylla var. aphylla, S. crassiramea, S. rigidicaulis and S. spiniflora; and Clade B grouping S. acanthoclada, S. aphylla var. divaricata, S. aphylla var. pendula and S. pachyrrhiza. The morphologically complex S. aphylla appears thus polyphyletic. Molecular dating and ancestral area reconstructions suggest that the Aphyllae clade started to diversify in the South American Transition Zone in the Late Pliocene. Based on these results, we hypothesize that this diversification occurred during the last period of Andean uplift with the aridification in South America. The ancestral character state reconstructions suggest that, in addition to the loss of leaves in adult plants, series Aphyllae evolved various morphological features, such as fastigiate, thickened or decumbent–subdecumbent branches during the colonization and establishment in different arid and semiarid lands in South America.

Mitochondrial phylogenomics of Hemiptera reveals adaptive innovations driving the diversification of true bugs.

Hemiptera, the largest non-holometabolous order of insects, represents approximately 7% of metazoan diversity. With extraordinary life histories and highly specialized morphological adaptations, hemipterans have exploited diverse habitats and food sources through approximately 300 Myr of evolution. To elucidate the phylogeny and evolutionary history of Hemiptera, we carried out the most comprehensive mitogenomics analysis on the richest taxon sampling to date covering all the suborders and infraorders, including 34 newly sequenced and 94 published mitogenomes. With optimized branch length and sequence heterogeneity, Bayesian analyses using a site-heterogeneous mixture model resolved the higher-level hemipteran phylogeny as (Sternorrhyncha, (Auchenorrhyncha, (Coleorrhyncha, Heteroptera))). Ancestral character state reconstruction and divergence time estimation suggest that the success of true bugs (Heteroptera) is probably due to angiosperm coevolution, but key adaptive innovations (e.g. prognathous mouthpart, predatory behaviour, and haemelytron) facilitated multiple independent shifts among diverse feeding habits and multiple independent colonizations of aquatic habitats.

The morphological and functional variability of pleon-holding mechanisms in selected Eubrachyura (Crustacea: Decapoda)

ABSTRACTEubrachyurans not only exhibit a crab-like body shape, consisting of a mostly wide cephalothorax and a thin, shield-like pleon folded underneath it, but reportedly hold the pleon in this position with the use of specialized holding structures. To further our knowledge of the variability of these holding structures, and investigate their functionality, male and female crabs from five primary freshwater crab and five selected thoracotreme families were examined with stereomicroscopy. Exemplary specimens of each group were documented photographically and with micro-computed tomography to illustrate the holding structures in situ. The results show a press-button mechanism of variable shape in freshwater crabs, where divergent structures possibly support pleon attachment of different effectiveness. The press-button is present in the majority of basal heterotreme lineages, and is therefore regarded as an ancestral character of the Heterotremata. In the Thoracotremata, multiple holding mechanisms are identified: press-button, clasping apparatus, overhanging edge, and increased setae cover on sternal plates, with the last system probably only effective in terrestrial species, due to wet adhesion effects. The ancestral character state in the Thoracotremata is debatable, but most likely also a functional press-button was present. Each holding mechanism requires different pleon movements to engage and disengage. Additionally, in contrast to the most common press-button mechanism, divergent holding mechanisms tend to be present, and likely functional, in adult males and females.

Molecular Tracing of the Geographical Origin of Human Immunodeficiency Virus Type 1 Infection and Patterns of Epidemic Spread Among Migrants Who Inject Drugs in Athens.

High numbers of human immunodeficiency virus type 1 (HIV-1) infections among people who inject drugs (PWID) have been diagnosed in Athens, Greece, since 2011. We aimed to trace the geographic origin of HIV-1 infection for migrants who inject drugs and to investigate whether transmissions occur more frequently among migrants than among Greek nationals. Multiple cross-sectional studies were pooled to assemble all persons diagnosed with HIV-1 in Greece between 1 January 2011 and 31 October 2014. Phylogenetic analyses used maximum likelihood estimation. The hypothesis of ethnic compartmentalization was tested by reconstructing ancestral states of characters at the tips using the criterion of parsimony over a set of bootstrap trees. Of 2274 persons, 38.4% were PWID. Phylogenetic analyses showed the existence of 4 major PWID-specific local transmission networks (LTNs): CRF14_BG (437 [58.6%]), CRF35_AD (139 [18.6%]), subtype B (116 [15.6%]), and subtype A (54 [7.2%]). Of 184 non-Greek PWID, 78.3% had been infected within the PWID-LTNs. For 173 (94.3%), the origin of their infection was assumed to be in Greece (postmigration). For PWID infected within LTNs, transmissions for subtype A and CRF14_BG occurred more frequently among migrants than would be expected by chance (phyloethnic study). Our analysis showed that the majority of infections among migrants occurred postmigration. The existence of significant transmission networking among migrants highlights that this population is a priority for HIV prevention. As molecular analysis can estimate the probable country of HIV infection, it can help to inform the design of public health strategies.

A Burmese amber fossil of <i>Radula</i> (Porellales, Jungermanniopsida) provides insights into the Cretaceous evolution of epiphytic lineages of leafy liverworts

Abstract. DNA-based divergence time estimates suggested major changes in the composition of epiphyte lineages of liverworts during the Cretaceous; however, evidence from the fossil record is scarce. We present the first Cretaceous fossil of the predominantly epiphytic leafy liverwort genus Radula in ca. 100 Myr old Burmese amber. The fossil's exquisite preservation allows first insights into the morphology of early crown group representatives of Radula occurring in gymnosperm-dominated forests. Ancestral character state reconstruction aligns the fossil with the crown group of Radula subg. Odontoradula; however, corresponding divergence time estimates using the software BEAST lead to unrealistically old age estimates. Alternatively, assignment of the fossil to the stem of subg. Odontoradula results in a stem age estimate of Radula of 227.8 Ma (95 % highest posterior density (HPD): 165.7–306.7) and a crown group estimate of 176.3 Ma (135.1–227.4), in agreement with analyses employing standard substitution rates (stem age 235.6 Ma (142.9–368.5), crown group age 183.8 Ma (109.9–289.1)). The fossil likely belongs to the stem lineage of Radula subg. Odontoradula. The fossil's modern morphology suggests that switches from gymnosperm to angiosperm phorophytes occurred without changes in plant body plans in epiphytic liverworts. The fossil provides evidence for striking morphological homoplasy in time. Even conservative node assignments of the fossil support older rather than younger age estimates of the Radula crown group, involving origins for most extant subgenera by the end of the Cretaceous and diversification of their crown groups in the Cenozoic.

Fossil evidence of eupolypod ferns in the mid-Cretaceous of Myanmar

Divergence time estimates based on DNA sequence variation of extant species suggest that eupolypod ferns were diverse already in the Cretaceous; however, fossil evidence to support this assumption remains exceedingly rare. Holttumopteris burmensis gen. et sp. nov., a fertile fern foliage fragment preserved in a piece of Albian–Cenomanian Burmese amber from Myanmar, is characterized by divided fertile leaves with catadromous, free lateral veins. Sporangia possess a vertical annulus interrupted by the sporangium stalk and contain monolete spores with a lophate perine. The sporangia occur clustered in discrete sori overarched by a laterally attached, reniform indusium. Reconstruction of ancestral character states suggests affinities of Holttumopteris to the Thelypteridaceae; however, several taxonomically relevant characters are not preserved. This discovery is nevertheless important because H. burmensis represents the first unequivocal fossil of a representative of the eupolypods from the middle Cretaceous.

Evolutionary Scenarios and Primate Natural History

Scenarios summarize evolutionary patterns and processes by interpreting organismal traits and their natural history correlates in a phylogenetic context. They are constructed by (1) describing phenotypes (including physiology and behavior), ideally with attention to formative roles of development, experience, and culture; (2) inferring homologies, homoplasies, ancestral character states, and their transformations with phylogenetic analyses; and (3) integrating those components with ecological and other ancillary data. At their best, evolutionary scenarios are factually dense narratives that entail no known falsehoods; their empirical and methodological shortcomings are transparent, they might be rejected based on new discoveries, and their potential ideological pitfalls are flagged for scrutiny. They are exemplified here by homoplastic foraging with percussive tools by humans, chimpanzees, capuchins, and macaques; homoplastic hunting with spears by humans and chimpanzees; and private experiences (e.g., sense of fairness, grief) among diverse animals, the homologous or homoplastic status of which often remains unexplored. Although scenarios are problematic when used to bolster political agendas, if constructed carefully and regarded skeptically, they can synthesize knowledge, inspire research, engender public understanding of evolution, enrich ethical debates, and provide a deeper historical context for conservation, including nature appreciation.

Discovery of a new Mesozoic species of the ancient genus Lepicerus (Coleoptera: Myxophaga: Lepiceridae), with implications for the systematic placement of all previously described extinct ‘lepiceroids’

†Lepicerus mumia Jałoszyński & Yamamoto, sp. nov. is described based on a Cenomanian Myanmar amber inclusion. This is the third known and the best preserved fossil of ‘lepiceroid’ myxophagan beetles, making it possible to carry out a more detailed comparative study than previously published findings. Detailed analyses of diagnostic characters given by previous authors for †Haplochelus (†Haplochelidae) and †Lepichelus (= Lepiceroides) (Lepiceridae) led to conclusions that some features might have been misinterpreted, others do not differ from those in Lepicerus, and the remaining morphological structures can be regarded as variable within one genus. Consequently, Haplochelus and Lepichelus are placed as junior synonyms of Lepicerus, and the synonymy between Haplochelidae and Lepiceridae, previously suggested by other authors, is maintained. Lepicerus mumia, although classified in the extant genus, shows an unusual, presumably ancestral myxophagan character state, a partly divided protarsus, with a distinctly demarcated, enlarged and curved tarsomere 1. The presence of a crust partly covering the body of L. mumia suggests a similar biology as that of two extant species that live near river banks among other semiaquatic arthropods.

Microanatomical diversification of the zona pellucida in aplochelioid killifishes.

This study investigates zona pellucida (ZP) ultrastructure in fertilized eggs of annual killifishes (suborder Aplocheiloidei), a group of highly specialized fishes that are able to survive desiccation for several weeks to months before they hatch. Little is known about ZP or chorionic ultrastructure sustaining these life-history modes, so scanning electron microscopy (SEM) was used to describe this trait in a large number of aplocheiloids with a focus on the family Rivulidae and the genus Hypsolebias. New images of ZP ultrastructure for 52 aplocheiloid species are provided, more than doubling the number characterized thus far. The evolution of chorionic structure within this group is studied using these new data. Characters were coded into a morphological matrix and optimized onto a consensus phylogeny to assess phylogenetic signal and reconstruct ancestral character states. Although ZP characters seem highly homoplastic and exhibit a large amount of structural convergence among lineages, aplocheiloid killifishes have evolved a number of unique structures associated with the chorion. Some annual species seem to have lost long filaments because eggs are deposited in the soil instead of being adhered to aquatic plants.

Cranial suture complexity in caviomorph rodents (Rodentia; Ctenohystrica).

Due to their flexibility, sutures are regions that experience greater strains than the surrounding rigid cranial bones. Cranial sutures differ in their degree of interdigitation or complexity. There is evidence indicating that a more convoluted suture better enables the absorption of high stresses coming from dynamic masticatory forces, and other functions. The Order Rodentia is an interesting clade to study this because of its taxa with diverse chewing modes. Due to repeated loading resulting from gnawing and grinding, energy absorption by the sutures might be a crucial factor in these mammals. Species within the infraorder Caviomorpha were chosen as a case study because of their ecomorphological and dietary diversity. This study compared five sutures from the rostrum and cranial vault across seven caviomorph families, and assessed their complexity by means of the relative length and fractal dimension. Across these rodents, cranial sutures are morphologically quite diverse. We found that the sutures connecting the rostrum with the vault were relatively more interdigitated than those in the cranial vault itself, especially premaxillofrontal sutures. Suture interdigitation was higher in species that display chisel-tooth digging and burrowing behaviors, especially in the families Ctenomyidae and Octodontidae, than those in families Dasyproctidae and Cuniculidae, which have more gracile masticatory systems. The reconstruction of the ancestral character state, on family and species phylogeny, points toward low suture interdigitation (i.e., low length ratio) as a likely ancestral state for interfrontal, premaxillofrontal and maxillofrontal sutures. Interspecific differences in suture morphology shown here might represent adaptations to different mechanical demands (i.e., soft vs. tough foods) or behaviors (e.g., chisel-tooth digging), which evolved in close association with the diverse environments occupied by caviomorph rodents.

Multiple innovations underpinned branching form diversification in mosses.

Summary Broad‐scale evolutionary comparisons have shown that branching forms arose by convergence in vascular plants and bryophytes, but the trajectory of branching form diversification in bryophytes is unclear. Mosses are the most species‐rich bryophyte lineage and two sub‐groups are circumscribed by alternative reproductive organ placements. In one, reproductive organs form apically, terminating growth of the primary shoot (gametophore) axis. In the other, reproductive organs develop on very short lateral branches. A switch from apical to lateral reproductive organ development is proposed to have primed branching form diversification.Moss gametophores have modular development and each module develops from a single apical cell. Here we define the architectures of 175 mosses by the number of module classes, branching patterns and the pattern in which similar modules repeat. Using ancestral character state reconstruction we identify two stages of architectural diversification.During a first stage there were sequential changes in the module repetition pattern, reproductive organ position, branching pattern and the number of module classes. During a second stage, vegetative changes occurred independently of reproductive fate.The results pinpoint the nature of developmental change priming branching form diversification in mosses and provide a framework for mechanistic studies of architectural diversification.

Phylogenetic analysis and evolution of morphological characters in Metrodorea and related species in Rutoideae (Rutaceae)

The aim of the study is to compare distinct morphological characters of Metrodorea (Rutaceae) with those of related genera, under a phylogenetic approach. Morphoanatomy of leaves was analyzed, and data concerning reproductive structures were compiled from the literature. We used available molecular sequences to recover a maximum likelihood tree, and we present a reconstruction of ancestral morphological character states made with the maximum likelihood criterion. The topologies obtained retrieve as synapomorphic characters of Metrodorea: aborted leaflets, vascularized intrapetiolar stipules, sinuous cell walls contour in the surface view of the epidermis, glandular trichomes on the adaxial and proximal region of the petiole, good development of areoles, muricate fruits, and valvate estivation of the corolla. The specific relationships within Metrodorea and between Metrodorea and other genera of Rutoideae are supported by characters as the location of crystals, type of adherence at leaf bases, and presence/absence of trichomes on the blade. The close relationship of Metrodorea and Raulinoa is strongly supported by several synapomorphies. Our data are mostly consistent with recent phylogenies, showing that the monophyly of Esenbeckia cannot be confirmed and that the clade Helietta-Balfourodendron (formerly included in another subfamily) is closely related to Raulinoa and Metrodorea (both part of Pilocarpinae) than to Pilocarpus itself. Possible relationships between these genera are discussed based on the morphological states found, and most of them supporting the hypothesis that Metrodorea is the sister group of Raulinoa.

Trophic evolution in African citharinoid fishes (Teleostei: Characiformes) and the origin of intraordinal pterygophagy

The African freshwater suborder Citharinoidei (Characiformes) includes 110 species that exhibit a diversity of feeding modes comparable to those characteristic of more speciose groups such its sister, the Characoidei (2000+ species) or the distantly related Cichlidae (1600+ species). Feeding habits of the Citharinoidei range from generalist omnivores to highly specialized feeding modes including ectoparasitic fin-eating, i.e. pterygophagy. We examine diet preference evolution in the Citharinoidei using newly inferred multi-gene-based hypotheses of phylogenetic relationships for representatives of 12 of the 15 genera in the suborder. Ancestral character state reconstructions onto our best tree indicate that the three most-generalist diets - pelophage/planktivore, omnivore and invertivore - are also the most primitive conditions within the Citharinoidei. The feeding mode of the most recent common ancestor of the Citharinoidei was characterized by high uncertainty. The more specialized feeding habits - herbivory, piscivory and pterygophagy - originated later in the Citharinoidei, likely from invertivore ancestors and possibly across a short time period. Highly specialized fin eaters (Belonophago, Phago and Eugnatichthys) share a common origin along with a strict piscivore (Mesoborus) and an invertivore (Microstomatichthyoborus). The largely piscivorous, but facultative fin eater, Ichthyborus is not exclusively related to them. Our results demonstrate that overall diet preference transitions in the Citharinoidei were rare events with very few reversals or parallelisms, and that evolutionary shifts in trophic ecology have not played a major role in intraordinal diversification. This situation contrasts with other groups in which dietary transitions have played key roles in species diversification.