Character Evolution Research Articles

TYPOLOGY OF FEMALE CHARACTERS IN ELEANOR PORTER’S NOVEL “THE ROAD TO UNDERSTANDING”

The article analyses the plot and imagery of Eleanor Porter’s novel “The Road to Understanding”. Using elements of the phenomenological approach to text analysis, comparative and contrastive analysis, and the methodology of cultural research, the author demonstrates the evolution of the main character’s image and comprehends the typology of women’s images in historical and social contexts. The authors outline the main features of the aesthetics, creative style and outlook of Eleanor Porter. The article proves that the author used irony, artistic detail, psychological narration, contraposition, portrait and speech characteristics of her heroes to reveal their female characters. The author analyses the correlation between the characters in the storyline, defines the importance of the role of minor characters in the main heroine’s character development. The special features of the concept “daughter” are shown as an embodiment of mental notions of spiritual beauty and family happiness. The value of the image-symbol of mother, its role in the composition and the artistic space of the work is revealed. Besides the comparative characteristic of characters in the work is given. In the process of work a search for new ways of analysis of artistic images in the artwork on the basis of interpretation and reader reception was carried out, as well as tradition and innovation in the work of the writer was revealed. The article proves artistic and moral value of Eleanor Porter’s heritage.

The absence of the evolutionary state of the Prokaryote would imply a polyphyletic origin of proteins and that LUCA, the ancestor of bacteria and that of archaea were progenotes

I analysed the similarity gradient observed in protein families – of phylogenetically deep fundamental traits - of bacteria and archaea, ranging from cases such as the core of the DNA replication apparatus where there is no sequence similarity between the proteins involved, to cases in which, as in the translation initiation factors, only some proteins involved would be homologs, to cases such as for aminoacyl-tRNA synthetases in which most of the proteins involved would be homologs. This pattern of similarity between bacteria and archaea would seem to be a very clear indication of a transitional evolutionary stage that preceded both the Last Bacterial Common Ancestor and the Last Archaeal Common Ancestor, i.e. progenotic stages. Indeed, this similarity pattern would seem to exemplify an ongoing transition as all the evolutionary phases would be represented in it. Instead, in the cellular stage it is expected that these evolutionary phases should have already been overcome, i.e. completed, and therefore no longer detectable. In fact, if we had really been in the presence of the prokaryotic stage then we should not have observed this similarity pattern in proteins involved in defining the ancestral characters of bacteria and archaea, as the completion of the different cellular structures should have required a very low number of proteins to be late evolved in lineages leading to bacteria and archaea. Indeed, the already reached state of the Prokaryote would have determined complete cellular structures therefore a total absence of proteins to evolve independently in the two main phyletic lineages and able to complete the evolution of a particular character already evidently in a definitive state, which, on the other hand, does not appear to have been the case. All this would have prevented the formation of this pattern of similarity which instead would appear to be real. In conclusion, the existence of this pattern of similarity observed in the families of homologous proteins of bacteria and archaea would imply the absence of the evolutionary stage of the Prokaryote and consequently a progenotic status to be assigned to the LUCA. Indeed, the LUCA stage would have been a stage of evolutionary transition because it is belatedly marked by the presence of all the different evolutionary phases, evidently more easily interpretable within the definition of progenote than that of genote precisely because they are inherent in an evolutionary transition and not to an evolution that has already been achieved. Finally, I discuss the importance of these arguments for the polyphyletic origin of proteins.

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

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

A PORTRAIT OF THE ARTIST AS A YOUNG MAN - AN INQUIRY INTO THE CHARACTER’S EVOLUTION

James Joyce, a prominent figure in the modernist literary movement, created the character Stephen Dedalus and revealed an example of how consciousness functions when one searches for direction in life. The chaotic nature of human consciousness is revealed through literary devices and techniques that prove the lifelikeness of the character and the way fiction represents not only the reality of a specific time but the reality of the multitude of meanings present within the content of fictional writing, namely literature. The aim of this paper is to investigate the evolution of the main character in the novel “A Portrait of The Artist as a Young Man” by James Joyce to reveal the way Stephen Dedalus builds his consciousness and finds the tone of his “voice”. The metaphor of the voice is to be considered the conceptual term used to describe the different phases of the character’s evolution and the way he succeeds in creating order among uncertainties, quests, and struggles. The term “voice” is used within the neuro-scientific frame of thought, as defined by Antonio Damasio as being “the representation of the conscious self” (Damasio 2010: 45). Both terms, namely consciousness and voice, lead to psychological content; thus, the nature of the analysis is psychological, literary, and symbolic as well. The mythological dimension is to be interpreted through symbols and their significance. The literary dimension revolves around literary techniques such as interior monologue and stream of consciousness, and the psychological frame includes elements that connect literature and psychology.

Monnina (Polygalaceae), a New World monophyletic genus full of contrasts

Abstract Endemic to the Neotropics, Monnina is the second largest genus of Polygalaceae, yet little is known about its phylogenetic history, biogeography, and morphological character evolution. To address these knowledge gaps, we conducted Bayesian and maximum likelihood (ML) analyses of nuclear ITS and plastid trnL–F regions to test the monophyly of Monnina s.l. We used this phylogenetic framework to (i) infer divergence time estimates of lineages within the genus and reconstruct their historical biogeography; (ii) reconstruct the evolution of morphological characters of putative ecological and evolutionary importance in Monnina; and (iii) test for correlations between our phylogenetic hypothesis and environmental data. Our results reveal that Monnina is monophyletic with an indehiscent, 1–2-seeded fruit as a synapomorphy for the genus. We identify six clades within Monnina based on our combined phylogenetic results: Clades A, B, and D are primarily distributed in southern and eastern South America, Clades C and E are primarily Central Andean, and Clade F is chiefly distributed in the Northern Andes and Central America. The ancestor of the Monnina stem lineage dispersed from Australia/Africa to South America during the late Eocene to early Oligocene. The divergences of major lineages within the genus began in the early Miocene. We inferred the most recent common ancestor of Monnina to be an herbaceous plant with one-seeded samaroid fruits. The origins of fleshy fruits and shrubby habits are phylogenetically correlated within Monnina, and their concerted convergent evolution may have promoted increased net diversification rates in the two most species-rich subclades of the genus.

Generation three and a half peacekeeping: Understanding the evolutionary character of African-led Peace Support Operations

ABSTRACT African-led Peace Support Operations (PSOs) were established to support the African peace and security architecture by developing integrated capacities for deployment in crises. However, since the deployment of the first African-led PSOs, there has also been the emergence of new types of African-led PSOs, such as the African Union Mission in Somalia; the Lake Chad Basin Commission Multinational Joint Task Force; the Joint Force for the Group of Five for the Sahel; the Southern African Development Community Mission in Mozambique; and the East African Community Force in Eastern Democratic Republic of Congo. The paper examines why African-led PSOs have emerged, arguing that these operations have allowed for increased African agency and shaped the African peace and security space. The paper finds that African-led PSO reflects a more regional and local-specific response in a declining era of new United Nations peacekeeping operations but has also resulted in an overreliance on force to solve the continent’s peace and security issues. Consequently, the paper arrives at a novel conceptualisation of African-led PSOs, positing that they represent generation three and a half of peacekeeping which focus on the effectiveness of force and the morality of using force to deal with insecurity and multifaceted crisis.

Evolutionary history, traits, and weediness inDigitaria(Poaceae: Panicoideae)

AbstractDigitaria is a large pantropical genus, which includes a number of economically problematic agricultural weeds. Difficulties in species identification and the circumscription of the genus have previously hindered progress in understanding its evolution and developing a stable classification. We investigate the evolutionary history of Digitaria by combining two approaches: (i) phylogenetic analyses of the currently available Sanger sequence data with previously unpublished sequences mainly from South East Asia, and (ii) newly generated genomic data from low-copy nuclear genes obtained using the Angiosperms353 probe kit, with new data mainly from Madagascar. Both methods gave congruent results. The combination of these results allowed us to confirm a new clade structure within Digitaria lineages, in partial agreement with the previous morphology-based classifications. We confirm that Digitaria as traditionally circumscribed is not monophyletic, comprising Anthephora, Baptorhachis, Chaetopoa, and Chlorocalymma. We present an inference of key character evolution with morphological support for clades with spikelets arranged in binate and ternate groups, with variable trichome morphology, spikelet length, and culm length. Geographical clade structure is also observed. No phylogenetic signal of weedy function was observed.

The Non-Dereliction in Evolution: Trophic Specialisation Drives Convergence in the Radiation of Red Devil Spiders (Araneae: Dysderidae) in the Canary Islands.

Natural selection plays a key role in deterministic evolution, as clearly illustrated by the multiple cases of repeated evolution of ecomorphological characters observed in adaptive radiations. Unlike most spiders, Dysdera species display a high variability of cheliceral morphologies, which has been suggested to reflect different levels of specialization to feed on isopods. In this study, we integrate geometric morphometrics and experimental trials with a fully resolved phylogeny of the highly diverse endemic species from the Canary Islands to 1) quantitatively delimit the different cheliceral morphotypes present in the archipelago, 2) test their association with trophic specialization, as reported for continental species, 3) reconstruct the evolution of these ecomorphs throughout the diversification of the group, 4) test the hypothesis of convergent evolution of the different morphotypes, and 5) examine whether specialization constitutes a case of evolutionary irreversibility in this group. We show the existence of 9 cheliceral morphotypes and uncovered their significance for trophic ecology. Further, we demonstrate that similar ecomorphs evolved multiple times in the archipelago, providing a novel study system to explain how convergent evolution and irreversibility due to specialization may be combined to shape phenotypic diversification in adaptive radiations.

Phylogeny of the tribe Riorajini with comments on the morphological character evolution within Rajiformes (Chondrichthyes, Batoidea)

AbstractThe tribe Riorajini is composed of four species, Rioraja agassizii, Atlantoraja castelnaui, A. cyclophora and A. platana, which are endemic to the inshore waters of the Southwestern Atlantic Ocean. Previous studies have hypothesized the monophyly of this group but interrelationships within its members remain not fully understood. In this study, we analysed quantitative as well as qualitative morphological characters related to external morphology, squamation, neurocranium, visceral arches, pectoral and pelvic girdles and claspers, totalizing 82 characters, representing a greater number than previously used in studies focused on rajiform systematics. 24 terminal taxa were included, and two different analyses were carried out testing Narcine brasiliensis and Pseudobatos horkelii as roots of the cladogram. The monophyly of Riorajini is supported by five quantitative and three discrete characters, including a caudal length smaller than disc length, sensory pores and mucous canals darkly pigmented on ventral surface of disc, and preorbital flange present. Clasper morphology was phylogenetically informative since it contributes to diagnose Rioraja in which an accessory terminal 2 cartilage is absent and to support a sister relationship between A. cyclophora + A. platana. Squamation, teeth and neurocranial morphology need to be thoroughly investigated in order to revise their terminology, propose and test hypotheses of homology and improve our knowledge on morphological variation regarding ontogeny and sexual dimorphism.

Plastomic data shed new light on the phylogeny, biogeography, and character evolution of the family Crassulaceae

AbstractCrassulaceae is a mid‐sized family of angiosperms, most species of which are herbaceous succulents, usually with 5‐merous flowers and one or two whorls of stamens. Although previous phylogenetic studies revealed seven major “clades” in Crassulaceae and greatly improved our understanding of the evolutionary history of the family, relationships among major clades are still contentious. In addition, the biogeographic origin and evolution of important morphological characters delimiting infrafamilial taxa have not been subject to formal biogeographic and character evolution analyses based on a well‐supported phylogeny backbone. In this study, we used plastomic data of 52 species, representing all major clades revealed in previous studies to reconstruct a robust phylogeny of Crassulaceae, based on which we unraveled the spatiotemporal framework of diversification of the family. We found that the family may originate in southern Africa and then dispersed to the Mediterranean, from there to eastern Asia, Macaronesia, and North America. The crown age of Crassulaceae was dated at ca. 63.93 million years ago, shortly after the Cretaceous–Paleogene (K‐Pg) boundary. We also traced the evolution of six important morphological characters previously used to delimit infrafamilial taxa and demonstrated widespread parallel and convergent evolution of both vegetative (life form and phyllotaxis) and floral characters (number of stamen whorls, petals free or fused, and flower merism). Our results provide a robust backbone phylogeny as a foundation for further investigations, and also some important new insights into biogeography and evolution of the family Crassulaceae.

Molecular phylogenetics of sessile Dolium sedentarium, a petalomonad euglenid.

The euglenids are a species-rich group of flagellates with varying modes of nutrition that can be found in diverse habitats. Phagotrophic members of this group gave rise to phototrophs and hold the key to understanding the evolution of euglenids as a whole, including the evolution of complex morphological characters like the euglenid pellicle. Yet to understand the evolution of these characters, a comprehensive sampling of molecular data is needed to correlate morphological and molecular data, and to estimate a basic phylogenetic backbone of the group. While the availability of SSU rDNA and, more recently, multigene data from phagotrophic euglenids has improved, several "orphan" taxa remain without any molecular data whatsoever. Dolium sedentarium is one such taxon: It is a rarely-observed phagotrophic euglenid that inhabits tropical benthic environments and is one of few known sessile euglenids. Based on morphological characters, it has been thought of as part of the earliest branch of euglenids, the Petalomonadida. We report the first molecular sequencing data for Dolium using single-cell transcriptomics, adding another small piece in the puzzle of euglenid evolution. Both SSU rDNA and multigene phylogenies confirm it as a solitary branch within Petalomonadida.

Insights into the morphology and evolution of orbicules in the Spermacoce clade (Spermacoceae-Rubiaceae) and implications for systematics

Abstract The orbicules are tiny structures of sporopollenin that occur on the interior wall of anthers in several groups of plants, and they are associated with pollen grains and tapetal cells. Although their function remains still unresolved, they have been widely used in systematics. Rubiaceae is one of the most studied families in this aspect. However, scarce information is available about the orbicules in the tribe Spermacoceae, especially in the Spermacoce clade. The main objectives of this work were to investigate the occurrence, general morphology, evolution, and systematic utility of the orbicular characters in the Spermacoce clade. We investigated the presence/absence, size, abundance, shape, and ornamentation of orbicules in 104 specimens from 84 species in the Spermacoce clade using scanning electron microscopy (SEM). On the basis of these features, the ancestral state reconstruction analyses were performed on a custom-made phylogeny by stochastic character mapping. The orbicules are present in 38 (44%) of 84 analysed species, corresponding to 13 genera (56%) of the Spermacoce clade. Seven genera have orbicules in all analysed species, eight genera lack orbicules in all analysed species, and six genera show both presence and absence of orbicules. The absence of orbicules was estimated to be the ancestral state for the Spermacoce clade and the presence of orbicules evolved several times independently within Clade B. The absence of orbicules in the common ancestor of the Spermacoce clade can be seen as one of the independent losses that occurred during the evolutionary history of the Rubiaceae.

An integrative view on the systematic position of the cupressophyteCephalotaxus

AbstractWe made an in‐depth review of historical studies of the cupressophyte conifer genusCephalotaxusSiebold & Zucc. with an emphasis on its systematic position. We suggest that the systematic position of the genus is better understood using an integrative approach, so the evolution of phenetic characters is discussed within the context of recent phylogenomics. We propose that the genus should be classified as a separate family Cephalotaxaceae belonging to the clade consisting of Cupressaceae, Cephalotaxaceae, and Taxaceae; the family Cephalotaxaceae is sister to the Taxaceae but not nested within the Taxaceae and is characterized by a unique set of characters including morphology, anatomy, embryology, and chemistry. The family Cephalotaxaceae shows transitional characters between the Cupressaceae and the Taxaceae; the family possesses female cones with a primary cone axis bearing 5–8 pairs of decussate bracts, which is similar to the typical female cones of the Cupressaceae, on the one hand, and may have given rise to the reduced female cone of the Taxaceae with one terminal ovule partially or completely enclosed in a fleshy aril. In parallel, the compound male cone of the Cephalotaxaceae evolved into the seemingly “simple” male cones of the Taxaceae by means of reduction, elimination, and fusion.

MODERN MILITARY PROSE: GENRE AND STYLE TYPOLOGY

The paper deals with the typology of modern Ukrainian military prose based on the analysis of relevant novels. According to the structural and functional features of the works, it is reasonable to classify them using division into high/mass/intermediate literature. The first type includes complex and multidimensional texts (with allusions, literary reminiscences, irony, parody, etc.), and their action takes place mainly in a mental sphere. Heroes are characterized by understanding of social processes and war realities (significant is the presence of two identities, “properly Ukrainian” and “Soviet”); they rethink their mistakes and transform in response to new challenges. These texts are mostly focused on the acquisition of individual subjectivity. They require a trained erudite reader who can discern the writer’s narratives behind their multi-layered nature. Mass literature includes works where action dominates, and such elements as shooting, chase, superman hero, excessive emotions, love triangle, etc. are widely used. The adventure genre, military detective, melodrama, and action novel are currently most fully represented. Intermediate prose may combine, on the one hand, topical social issues, character evolution, logical and consistent presentation of events (as a feature of “high literature”) with a simplified plot and images, the lack of attention to the psychology of the characters, and various means of mass literature. Contemporary Ukrainian writers have managed to create a large-scale cultural discourse, offering works that cover the problem of war, its causes and consequences, and the topic of a person at war from different perspectives and contexts.

MORPHO-PEDOLOGY VARIATIONS IN A PRE-SAHARAN REGION CASE ZIBAN SOILS

During this study, we tried to highlight the main lines of the spatial evolution of the morpho-pedological characters of certain soils, on a sequence established from the geomorphological map in the region. Biskra is an Oasis located in the North-Eastern Sahara of Algeria. After a long prospection, we have established two sequences of soils which have different morphological characteristics. These variations are mainly due to the geomorphological position of the terrain and the favorable hydrological aspect in the first sequence to the formation of gypsum soils compared to the second.

Taxonomic contributions to Hapalidiales (Corallinophycidae, Rhodophyta): Boreolithothamnion gen. nov., Lithothamnion redefined and with three new species and Roseolithon with new combinations.

Phylogenetic analyses of rbcL gene sequences and of concatenated rbcL, psbA, and nuclear SSU rRNA gene sequences resolved the generitype of Lithothamnion, L. muelleri, in a clade with three other southern Australian species, L. kraftii sp. nov., L. saundersii sp. nov., and L. woelkerlingii sp. nov. Cold water boreal species currently classified in Lithothamnion and whose type specimens have been sequenced are transferred to Boreolithothamnion gen. nov., with B. glaciale comb. nov. as the generitype. The other species are B. giganteum comb. nov., B. phymatodeum comb. nov., and B. sonderi comb. nov., whose type specimens are newly sequenced, and B. lemoineae comb. nov., B. soriferum comb. nov., and B. tophiforme comb. nov., whose type specimens were already sequenced. Based on rbcL sequences from the type specimens of Lithothamnion crispatum, L. indicum, and L. superpositum, each is recognized as a distinct species and transferred to the recently described Roseolithon as R. crispatum comb. nov., R. indicum comb. nov., and R. superpositum com. nov., respectively. To correctly assign species to these three genera based only on morpho-anatomy, specimens must have multiporate conceptacles and some epithallial cells with flared walls. The discussion provides examples demonstrating that only with phylogenetic analyses of DNA sequences can the evolution of morpho-anatomical characters of non-geniculate corallines be understood and applied at the correct taxonomic rank. Finally, phylogenetic analyses of DNA sequences support recognition of the Hapalidiales as a distinct order characterized by having multiporate tetra/bisporangial conceptacles, and not as a suborder of Corallinales whose tetra/bisporangial conceptacles are uniporate.

Palynological and macroscopic characters evidence infer the evolutionary history and insight into pollination adaptation in Impatiens (Balsaminaceae)

AbstractThe genus Impatiens comprises more than 1000 species and displays enormous morphological diversity. As a taxon that is notoriously difficult to classify morphologically, the taxonomic significance of pollen morphology has been discussed in previous studies. However, there was only limited study on the evolution of palynological and macroscopic character in this genus before. In this paper, we observed pollen of 143 species of Impatiens, including all representatives of infrageneric groups and discussed their taxonomic value. Meanwhile, based on the reconstructed phylogenetic framework, we selected five pollen traits, and nine macroscopic traits including both vegetative and reproductive organs for evolutionary analysis. The character reconstruction showed that the morphology and number of aperture, granules in lumina, seed number of per capsule, number of sepals, lateral united petals connate, spur length, and phyllotaxy could be identified as unambiguous, while other characters were ambiguous in the last common ancestor of Impatiens. The results of comprehensively macroscopic characters and palynological evidence suggest that these traits experienced a complicated evolution, exhibit high levels of instability and variability and occur character multiple reversals in the evolutionary processes of Impatiens due to different selection pressures. In addition, we found that there is an evident mosaic evolution in this genus. Pollen and macroscopic character are conserved with a slower evolutionary rate in the I. subgen. Clavicarpa, however, these characters showed higher diversity with a rapid rate of evolution in the I. subgen. Impatiens. Furthermore, the relationship between floral morphology and pollinators was discussed.

EvoPhylo: An r package for pre‐ and postprocessing of morphological data from relaxed clock Bayesian phylogenetics

Abstract Relaxed clock Bayesian evolutionary inference (BEI) enables the co‐estimation of phylogenetic trees and evolutionary parameters associated with models of character and lineage evolution. Fast advances in new model developments over the past decade have boosted BEI as a major macroevolutionary analytical framework using morphological and/or molecular data across vastly different study systems. However, there is limited availability of bioinformatic tools to pre‐ and postprocess data from BEI, such as identifying morphological data partitions, or statistically testing and creating publication quality plots of evolutionary hypotheses. Here, we introduce EvoPhylo, an r package to perform automated morphological character partitioning and analyse macroevolutionary parameter from relaxed clock (time‐calibrated) BEI outputted by the programs Mr.Bayes and BEAST2. These include rates of evolution and mode of selection for each character partition, diversification rate parameters, and handling fossil‐only posterior trees. We present the theoretical background behind EvoPhylo's functions and analytical tools for evolutionary hypothesis testing, its potential uses, and interpretation of its results with a series of vignettes and links to a step‐by‐step tutorial using examples from two empirical datasets. EvoPhylo will facilitate the use of Bayesian relaxed clocks as a tool for macroevolutionary inference across a wide range of users and fields of research, especially those that make usage of morphological datasets, from paleontological to total evidence dating analyses.

Character evolution of modern eupolypods supports the assignment of the fossil fern Cretacifilix fungiformis to Dryopteridaceae

The fossil fern species Cretacifilix fungiformis from mid-Cretaceous Kachin amber from Myanmar was previously assigned to eupolypod ferns for having sporangia with a vertical annulus and sporangial stalks 2–3 cells thick, along with monolete spores with a distinct perine. However, due to the insufficient documentation of character states and the unavailability of the holotype, the proposed affiliation of this fossil to derived families of eupolypods required further study. The recent discovery of additional leaf fragments of C. fungiformis from the same amber deposit enabled us to re-evaluate the relationships of this fossil taxon. Newly documented morphological evidence corroborated its placement within the eupolypod ferns. This was achieved by evaluating identifiable features present in the fossil species through an ancestral character state reconstruction utilizing a comprehensive, time-calibrated phylogeny of eupolypod ferns. Our character state reconstruction of extant eupolypod families and genera recovered morphological similarity of Cretacifilix to the closely related genera Arachniodes and Dryopteris, supporting an assignment of this fossil genus to the species-rich family Dryopteridaceae within the Polypodiineae.

Plastome evolution and phylogenomics of Trichosporeae (Gesneriaceae) with its morphological characters appraisal.

Trichosporeae is the largest and most taxonomically difficult tribe of Gesneriaceae due to its diverse morphology. Previous studies have not clarified the phylogenetic relationships within this tribe on several DNA markers, including the generic relationships within subtribes. Recently, plastid phylogenomics have been successfully employed to resolve the phylogenetic relationships at different taxonomic levels. In this study, plastid phylogenomics were used to explore the relationships within Trichosporeae. Eleven plastomes of Hemiboea were newly reported. Comparative analyses, phylogeny and morphological character evolution within Trichosporeae were conducted on 79 species representing seven subtribes. The Hemiboea plastomes range from 152,742 bp to 153,695 bp in length. Within Trichosporeae, the sampled plastomes range from 152,196 bp to 156,614 bp and GC content from 37.2% to 37.8%. A total of 121-133 genes were annotated in each species, including 80-91 protein-coding genes, 34-37 tRNA genes, and 8 rRNA genes. The contraction and expansion of IR borders were not detected, and gene rearrangements and inversions did not occur. The 13 hypervariable regions were proposed as the potential molecular markers for species identification. A total of 24,299 SNPs and 3,378 indels were inferred, and most of the SNPs were functionally missense and silent variations. There were 1968 SSRs, 2055 tandem repeats and 2802 dispersed repeats. The RSCU and ENC values indicated that the codon usage pattern was conserved in Trichosporeae. Both the phylogenetic frameworks based on the whole plastome and 80 CDSs were basically concordant. The sister relationships between Loxocarpinae and Didymocarpinae were confirmed, and Oreocharis was a sister group of Hemiboea with high support. The morphological characters showed a complex evolutionary pattern of Trichosporeae. Our findings may contribute to future research on genetic diversity, morphological evolutionary patterns, and conservation of the tribe Trichosporeae.