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1.
Phylogenetic studies addressing relationships among chrysidid wasps have been limited. There are no hypotheses proposed for the Neotropical lineages of Chrysidini other than the classic cladogram published in the 1990s by Kimsey and Bohart. Herein we present a cladistic analysis based on 64 morphological characters coded for 54 species of Chrysidini, 32 of them being Ipsiura and 22 representing Caenochrysis, Chrysis, Exochrysis, Gaullea, Neochrysis, and Pleurochrysis. The species of Ipsiura were recovered as monophyletic and as the sister clade of Neochrysis in all most parsimonious trees. We discuss the high plasticity of some morphological characters as evidenced by their high homoplasy in the phylogenetic results, and we clarify the main morphological changes inferred on the phylogenetic tree for this genus. The effects of the inferred homoplasy were evaluated under an implied weighting cladistic analysis, and from a probabilistic perspective with Bayesian inference. Those alternative strategies did not alter the general conclusions about the monophyly of Ipsiura or the generic relationships in Chrysidini (changes were noticed in the species‐level relationships within certain parts of Ipsiura, where low branch support was common across all approaches). Among the species groups proposed by Linsenmaier (1985), only the marginalis group was recovered as monophyletic. We also evaluated the convoluted biogeographic history of the group. The resulting historical reconstructions indicate a complicated scenario of diversification of these wasps in the Neotropics, and a close association with forested biomes is discussed.  相似文献   

2.
Cuckoo wasps (Hymenoptera: Chrysididae) are a species‐rich family of obligate brood parasites (i.e. parasitoids and kleptoparasites) whose hosts range from sawflies, wasps and bees, to walking sticks and moths. Their brood parasitic lifestyle has led to the evolution of fascinating adaptations, including chemical mimicry of host odours by some species. Long‐term nomenclatural stability of the higher taxonomic units (e.g. genera, tribes, and subfamilies) in this family and a thorough understanding of the family's evolutionary history critically depend on a robust phylogeny of cuckoo wasps. Here we present the results from phylogenetically analysing ten nuclear‐encoded genes and one mitochondrial gene, all protein‐coding, in a total of 186 different species of cuckoo wasps representing most major cuckoo wasp lineages. The compiled data matrix comprised 4946 coding nucleotide sites and was phylogenetically analysed using classical maximum‐likelihood and Bayesian inference methods. The results of our phylogenetic analyses are mostly consistent with earlier ideas on the phylogenetic relationships of the cuckoo wasps' subfamilies and tribes, but cast doubts on the hitherto hypothesized phylogenetic position of the subfamily Amiseginae. However, the molecular data are not fully conclusive in this respect due to low branch support values at deep nodes. In contrast, our phylogenetic estimates clearly indicate that the current systematics of cuckoo wasps at the genus level is artificial. Several of the currently recognized genera are para‐ or polyphyletic (e.g. Cephaloparnops, Chrysis, Chrysura, Euchroeus, Hedychridium, Praestochrysis, Pseudochrysis, Spintharina, and Spinolia). At the same time, our data support the validity of the genus Colpopyga, previously synonymized with Hedychridium. We discuss possible solutions for how to resolve the current shortcomings in the systematics of cuckoo wasp genera and decided to grant Prospinolia the status of a valid genus (Prospinolia stat.n. ) and transferring Spinolia theresae [du Buysson 1900] from Spinolia to Prospinolia (Prospinolia theresae stat.restit. ). We discuss the implications of our phylogenetic inferences for understanding the evolution of host associations in this group. The results of our study not only shed new light on the evolutionary history of cuckoo wasps, but also set the basis for future phylogenomic investigations on this captivating group of wasps by guiding taxonomic sampling efforts and the design of probes for target DNA enrichment approaches.  相似文献   

3.
Chromosome number and morphology are relevant aspects of genomic organization of eukaryotes and are considered key components to comprehend evolutionary mechanisms and karyotypic differentiation. Social wasps belonging to the tribe Epiponini show complex social characteristics that make these insects interesting models for genetic and evolutionary studies. However, there is a paucity of genetic information in social wasps as a whole. Aiming to investigate the process of chromosomal evolution of the social Epiponini wasps, chromosomes of ten species of this group were analyzed using Giemsa and fluorochrome staining techniques and fluorescence in situ hybridization in two species. In this study a high variation in the chromosome number and morphology was found and the previous range of chromosome number of Epiponini was broadened from n = 8–32 to n = 5–33. We also suggest that chromosomal segments with high GC content must have had a key role in the karyotype diversification of these wasps. Moreover, based on a phylogenetic background we find evidence of a main role of chromosomal fusion in the occurrence of gradual decrease of chromosome number in Epiponini during its chromosomal evolutionary history.  相似文献   

4.
Microbial diseases are important selective agents in social insects and one major defense mechanism is the secretion of cuticular antimicrobial compounds. We hypothesized that given differences in group size, social complexity, and nest type the secretions of these antimicrobials will be under different selective pressures. To test this we extracted secretions from nine wasp species of varying social complexity and nesting habits and assayed their antimicrobial compounds against cultures of Staphylococcus aureus. These data were then combined with phylogenetic data to provide an evolutionary context. Social species showed significantly higher (18x) antimicrobial activity than solitary species and species with paper nests showed significantly higher (11x) antimicrobial activity than those which excavated burrows. Mud-nest species showed no antimicrobial activity. Solitary, burrow-provisioning wasps diverged at more basal nodes of the phylogenetic trees, while social wasps diverged from the most recent nodes. These data suggest that antimicrobial defences may have evolved in response to ground-dwelling pathogens but the most important variable leading to increased antimicrobial strength was increase in group size and social complexity.  相似文献   

5.
The subfamily Microgastrinae is a highly diversified group of parasitoid wasps that attacks all of the different groups of Lepidoptera. We explore here the phylogenetic signal in three gene (mitochondrial COI and 16S, and nuclear 28S) fragments as an assessment of their utility in resolving generic relationships within this species-rich insect group. These genes were chosen because their level of sequence divergence is thought to be appropriate for this study and because they have resolved relationships among other braconid wasps at similar taxonomic levels. True phylogenetic signal, as opposed to random signal or noise, was detected in the 16S and 28S data sets. Phylogenetic analyses conducted on each microgastrine data set, however, have all resulted in poorly resolved trees, with most clades being supported by low bootstrap values. The phylogenetic signal, if present, is therefore concentrated on a few well-supported clades. Some rapidly evolving sites may be too saturated to be phylogenetically useful. Nonetheless, the sequence data (nearly 2300 nucleotides) used here appear to exhibit the appropriate level of variation, theoretically, to resolve the relationships studied. Moreover, the clades that are well supported by the data are usually supported by more than one data set and represent different levels of sequence divergence. We suggest that the lack of phylogenetic signal observed is an indication of the presence of many short internal branches on the phylogeny being estimated, which in turn might be the result of a rapid diversification of the taxa examined. Relative specialization of diet, which is typically associated with parasitic behavior, is believed to result in high radiation rates, which may have been especially high in microgastrine wasps because of the great diversity of their lepidopteran hosts. This hypothesis of a rapid diversification caused by an abundance of host species remains speculative and more data will be needed to test it further.  相似文献   

6.

Background

Host-parasite interactions are among the most important biotic relationships. Host species should evolve mechanisms to detect their enemies and employ appropriate counterstrategies. Parasites, in turn, should evolve mechanisms to evade detection and thus maximize their success. Females of the European beewolf (Philanthus triangulum, Hymenoptera, Crabronidae) hunt exclusively honeybee workers as food for their progeny. The brood cells containing the paralyzed bees are severely threatened by a highly specialized cuckoo wasp (Hedychrum rutilans, Hymenoptera, Chrysididae). Female cuckoo wasps enter beewolf nests to oviposit on paralyzed bees that are temporarily couched in the nest burrow. The cuckoo wasp larva kills the beewolf larva and feeds on it and the bees. Here, we investigated whether H. rutilans evades detection by its host. Since chemical senses are most important in the dark nest, we hypothesized that the cuckoo wasp might employ chemical camouflage.

Results

Field observations suggest that cuckoo wasps are attacked by beewolves in front of their nest, most probably after being recognized visually. In contrast, beewolves seem not to detect signs of the presence of these parasitoids neither when these had visited the nest nor when directly encountered in the dark nest burrow. In a recognition bioassay in observation cages, beewolf females responded significantly less frequently to filter paper discs treated with a cuticular extract from H. rutilans females, than to filter paper discs treated with an extract from another cuckoo wasp species (Chrysis viridula). The behavior to paper discs treated with a cuticular extract from H. rutilans females did not differ significantly from the behavior towards filter paper discs treated with the solvent only. We hypothesized that cuckoo wasps either mimic the chemistry of their beewolf host or their host's prey. We tested this hypothesis using GC-MS analyses of the cuticles of male and female beewolves, cuckoo wasps, and honeybee workers. Cuticle extracts of Hedychrum nobile (Hymenoptera: Chrysididae) and Cerceris arenaria (Hymenoptera: Crabronidae) were used as outgroups. There was little congruence with regard to cuticular compounds between H. rutilans females and honeybees as well as females of C. arenaria and H. nobile. However, there was a considerable similarity between beewolf females and H. rutilans females. Beewolf females show a striking dimorphism regarding their cuticular hydrocarbons with one morph having (Z)-9-C25:1 and the other morph having (Z)-9-C27:1 as the major component. H. rutilans females were more similar to the morph having (Z)-9-C27:1 as the main component.

Conclusion

We conclude that H. rutilans females closely mimic the composition of cuticular compounds of their host species P. triangulum. The occurrence of isomeric forms of certain compounds on the cuticles of the cuckoo wasps but their absence on beewolf females suggests that cuckoo wasps synthesize the cuticular compounds rather than sequester them from their host. Thus, the behavioral data and the chemical analysis provide evidence that a specialized cuckoo wasp exhibits chemical mimicry of the odor of its host. This probably allows the cuckoo wasp to enter the nest with a reduced risk of being detected by olfaction and without leaving traitorous chemical traces.  相似文献   

7.
Advances in potter wasp systematics have been achieved recently, with classificatory changes resulting from analyses based upon large scale molecular datasets. For the Neotropics, recent hypotheses point to the occurrence of an exclusive clade recognized within the tribe Eumenini. In this group, several contributions regarding taxonomy and systematics have been proposed in the last five years, including the genus Alphamenes. This taxon contains seven described species whose distribution is exclusively Neotropical. Females are morphologically homogeneous, and characters related to copulatory organs are useful in male diagnosis. This contribution forms the first phylogenetic approach to include all species of Alphamenes, hence the first to strongly test for group monophyly. Our cladistic results recovered Alphamenes as a monophyletic group supported by male genital features. Relationships among included species also rely upon genitalic characters, highlighting the importance of these attributes for eumenine systematics. Recent phylogenetic investigations applied to the Neotropical fauna of potter wasps represent desirable advancements towards a natural classification for the group.  相似文献   

8.
Most species in Melastomataceae have poricidal anthers related to specialised bee buzz‐pollination, while some have anthers with large openings associated to non‐bee pollination systems. We tracked the evolution of anther morphology and seed number on the Miconieae phylogenetic tree to understand the evolutionary shifts in such pollination systems. Anther morphometric data and seed number were recorded for 54 taxa. Pollinators (bees, flies, wasps) were recorded for 20 available species. Ancestral state reconstruction was made using Maximum Likelihood from nrITS sequences. We used phylogenetic eigenvector regressions to estimate phylogenetic signal and the adaptive component for these traits. Species pollinated by bees or bees and wasps tend to have smaller pores and fruits with more seeds. Species pollinated by flies or flies and bees and/or wasps tend to have larger pores and fruits with less seeds. Independent evolution occurred three times for anthers with large pores and twice for fruits with few seeds. We detected a phylogenetic signal in both traits, and negative correlated evolution between them. In actinomorphic small‐flowered Miconieae, changes in anther morphology can be related to generalisation in the pollination system incorporating flies and wasps as pollinators and lessening the importance of buzzing bees in such process. Differences in pollen removal and deposition may explain differences in anther morphology and seed number in Miconieae.  相似文献   

9.
Morphometric data for 30 species of swarming wasps (Vespidae: Polistinae: Epiponini) are presented, representing all currently recognized genera. Data are coded according to whether females that were shown by dissection to be egglayers are larger, similar, or smaller for each dimension than non-egglayers. These data are analysed in a phylogenetic framework with primitively social Polistes and Mischocyttarus as outgroups. Representative measurements are illustrated to show that most genera of Epiponini appear to have ancestry in a lineage that has no queen caste comparable with either the primitively social outgroups, or the more derived species of the tribe. This analysis indicates that a conspiracy of workers that operates without a queen characterizes the societies of many Epiponini, or their recent ancestors.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 509–522.  相似文献   

10.
Chalcidoid wasps represent one of the most speciose superfamilies of animals known, with ca. 23,000 species described of which many are parasitoids. They are extremely diverse in body size, morphology and, among the parasitoids, insect hosts. Parasitic chalcidoids utilise a range of behavioural adaptations to facilitate exploitation of their diverse insect hosts, but how host use might influence the evolution of body size and morphology is not known in this group. We used a phylogenetic comparative analysis of 126 chalcidoid species to examine whether body size and antennal size showed evolutionary correlations with aspects of host use, including host breadth (specificity), host identity (orders of insects parasitized) and number of plant associates. Both morphological features and identity of exploited host orders show strong phylogenetic signal, but host breadth does not. Larger body size in these wasps was weakly associated with few plant genera, and with more specialised host use, and chalcidoid wasps that parasitize coleopteran hosts tend to be larger. Intriguingly, chalcidoid wasps that parasitize hemipteran hosts are both smaller in size in the case of those parasitizing the suborder Sternorrhyncha and have relatively larger antennae, particularly in those that parasitize other hemipteran suborders. These results suggest there are adaptations in chalcidoid wasps that are specifically associated with host detection and exploitation.  相似文献   

11.
  1. Brood parasitism is a breeding strategy adopted by many species of cuckoos across the world. This breeding strategy influences the evolution of life histories of brood parasite species.
  2. In this study, we tested whether the degree on diet specialization is related to the breeding strategy in cuckoo species, by comparing brood parasite and nonparasite species. We measured the gradient of diet specialization of cuckoos, by calculating the Gini coefficient, an index of inequality, on the multiple traits describing the diet of species. The Gini coefficient is a measure of statistical dispersion on a scale between 0 and 1, reflecting a gradient from low to high specialization, respectively. First, we tested the strength of the phylogenetic signal of diet specialization index among cuckoo species worldwide. Then, we ran phylogenetic generalized least square (PGLS) models to compare diet specialization, distribution range, and body mass of parasitic and nonparasitic cuckoo species, considering the phylogenetic signal of data.
  3. After adjusting for the phylogenetic signal of the data and considering both, species distribution range and species body mass, brood parasitic cuckoos were characterized by higher diet specialization than nonbrood parasitic species. Brood parasitic species were also characterized by a larger breeding distribution range than nonparasitic species.
  4. The findings of this study provide an additional understanding of the cuckoos’ ecology, relating diet and breeding strategies, information that could be important in conservation ecology.
  相似文献   

12.
The first comprehensive phylogenetic study of the wasp tribe Cryptini (Hymenoptera, Ichneumonidae, Cryptinae) is presented, based on 109 morphological characters and molecular data from seven loci. The dataset includes 370 species, 308 of which are from Cryptini, covering 182 of its 250 genera. Results from parsimony and likelihood analyses are generally congruent. The topology has several implications for ichneumonid higher‐level classification. Previous definitions of the Ichneumoniformes clade are supported, though newly including the Microleptinae. The cryptine subtribe Ateleutina is consistently recovered outside of the Cryptini clade and should be treated as a separate subfamily, Ateleutinae stat.n. The tribe Phygadeuontini is shown to be polyphyletic: while most of the sampled taxa were recovered in a single clade, many of its members are more closely related to the Ichneumoninae, Ateleutinae or Cryptini. Pending a more detailed study, the group should be treated as a separate subfamily, Phygadeuontinae stat. rev . The former Hemigastrini are recovered as largely monophyletic but with important exceptions. Hemigaster Brullé is recovered as part of the Phygadeuontini and is transferred to that group. Echthrus Gravenhorst is consistently recovered as part of Cryptini, rendering Aptesini as the correct name for the tribe. The subfamily Cryptinae should be restricted to the tribes Aptesini and Cryptini. Within Cryptini, the results show little support for the current subtribal classification, with most subtribes recovered as polyphyletic. A number of relatively stable clades are identified and discussed, but the relationships among them are weakly supported. Most of these clades are morphologically heterogeneous and building a subtribal classification based on them would be ineffectual; they are therefore treated under the informal designation of genus groups. The results highlight the ubiquity of morphological homoplasy in Cryptini, and provide a framework from which to address further systematic and evolutionary questions on this hyperdiverse group of parasitic wasps.  相似文献   

13.
The Ampulicidae are one of the most basal groups within the apoid wasps, the paraphyletic assemblage of wasps previously known as Sphecidae. The morphology and taxonomy of the Ampulicidae are poorly studied, and there is no strict cladistic approach on their phylogeny yet. Here we assemble morphological characters of phylogenetic significance and submit them to parsimony analyses using modern cladistic methods. This produces a well-resolved and firmly supported phylogenetic hypothesis on the generic relationships within the group. Based on our phylogenetic results a revised classification is provided, subdividing the Ampulicidae into two monophyletic subfamilies, Ampulicinae ( Ampulex and Trirogma ) and Dolichurinae, the latter comprising two monophyletic tribes, Dolichurini ( Dolichurus and Paradolichurus ) and Aphelotomini, new tribe ( Aphelotoma and Riekefella ). Based on the resulting cladogram, the historical biogeography and the fossil record of Ampulicidae are briefly discussed.
© The Willi Hennig Society 2009.  相似文献   

14.
Abstract  Allodapine bees present particular problems for taxonomy due to a high level of morphological conservatism in adults, even between genera. However, this tribe of bees also presents a unique opportunity to explore the evolution of social parasitism because of the comparatively large number of origins of socially parasitic species. Morphological differences presented here, along with DNA sequence data and molecular phylogenetic analyses, indicate a much larger number of Australian social parasite species in the genus Inquilina than previously anticipated, and suggest that the final number of socially parasitic species may be considerable. We describe five new species and present sequence data that will help elucidate the delineation of further new species. Inquilina provides a unique opportunity to study the evolution of social parasitism in social insects, but further studies will need to encompass both population genetic and phylogenetic approaches.  相似文献   

15.
盔唇瓢虫族昆虫是瓢虫科中一类重要的捕食性天敌,主要捕食介壳虫,是该类害虫的重要生物防治天敌。盔唇瓢虫族昆虫研究历史悠久,目前世界盔唇瓢虫族包括22属超过280种,其中有9属42种分布在中国。该族许多属建立时间早,鉴别特征的原始描述不够详细,导致属间界定模糊,属的地位时常受到质疑且变动频繁。一些种类丰富,形态多变、且分布广泛的属如盔唇瓢虫属、光缘瓢虫属仍缺少世界性的订正研究。随着瓢虫科分子系统发育研究的开展,瓢虫科经典分类系统受到了较大的挑战,而盔唇瓢虫亚科已被证实为非单系起源的类群。目前,将盔唇瓢虫族归入瓢虫亚科得到绝大多数瓢虫科分子系统发育研究的支持,但与其它类群的亲缘关系仍存争议。尽管已有研究对盔唇瓢虫族现存所有属的系统发育关系进行了分析,但部分类群缺少分子数据,导致这些类群在该族系统发育关系中的位置并不明确。本文回顾了国内外盔唇瓢虫族的分类历史,记述了目前该族在瓢虫科的分类地位以及族下系统发育关系进展,最后对盔唇瓢虫族分类研究存在的问题及未来研究进行了展望。  相似文献   

16.
Classifications of Theaceae have usually placed the endangered monotypic genus Apterosperma in tribe Schimeae (x=18), whereas recent molecular phylogenetic evidence supports its transfer to tribe Theeae (x=15). Molecular data have not resolved the phylogenetic position of Apterosperma within Theeae. We investigated the chromosome number and karyotype of Apterosperma in the context of molecular and morphological phylogenetic evidence to provide further insight into the placement of Apterosperma within Theaceae. The chromosome number and karyotype was found to be 2n = 30 = 26m + 4sm, consistent with the transfer of Apterosperma to tribe Theeae. When the chromosome data were incorporated into a data set of 46 other nonmolecular characters, Apterosperma was placed as the first-diverging lineage within the clade comprising tribe Theeae. This supports its placement based on molecular data. The low intrachromosomal asymmetry (type 1A) of Apterosperma, presumably ancestral for the family, is also consistent with this placement. Character optimization strongly supports a base chromosome number of x=15 for tribe Theeae. Because of variable and sometimes conflicting chromosome count reports of species in tribes Schimeae and Stewartieae, the base chromosome number of Theaceae could be either x=15 or 17.  相似文献   

17.
Monophyly and intratribal relationships of the tribe Trypetini were tested using mitochondrial 16S ribosomal RNA gene from 16 species of Trypetini, six tephritid outgroups, and two non-tephritid outgroups. The number of aligned sites was 1279 bp, but 1165 bp were used for analysis after excluding sites with missing data or gaps. Among these 1165 sites, 447 sites were variable and 305 were informative for parsimony analysis. Phylogenetic information was extracted from this data set using neighbor-joining and maximum parsimony methods and compared to a phylogenetic hypothesis proposed from the morphological literature. My molecular data suggest: (1) monophyly of the tribe Trypetini; (2) monophyly of the Chetostoma group; (3) monophyly of the Trypeta group; and (4) non-monophyletic nature of the previous concept of Hemilea. I suggest that more sensitive genetic markers with less A+T bias are needed to bring about finer resolution within the tribe Trypetini, and additional tephritid lineages should be sampled to identify the sister-group relationship of the tribe Trypetini.  相似文献   

18.
Social insect queens reproduce while workers generally do not. Queens may also have other behavioural roles in the colony. In small, independent-founding colonies of social wasps, the dominant queen physically enforces her interests over those of the workers and serves as a pacemaker of the colony, stimulating workers to forage and engage in other tasks. By contrast, in large-colony, swarm-founding wasps, the collective interests of the workers are fulfilled in sex allocation and production of males, whether or not they coincide with the interests of the queens. The behavioural role of the queens in such species has not been extensively studied. We investigated the role of the queens both in regulating worker activity and in reducing the numbers of reproductively active queens in the swarm-founding epiponine wasp Parachartergus colobopterus. We found no evidence that queens regulate worker activity, as they were rarely involved in any interactions. Worker activity may be self-organized, without centralized active control by anyone. Furthermore, we found no evidence that the reduction in queen number characteristic of this tribe of wasps occurs in response to aggression among queens. The reduction in queen number may be a result of worker treatment of queens, although worker discrimination against some queens was not obvious in our data. i Copyright 2000 The Association for the Study of Animal Behaviour.  相似文献   

19.
The evolution of parental care strategies in aculeate (stinging) wasps and bees has been much studied from a functional perspective, but relatively little phylogenetic information is available to place this in a rigorous historical context, especially at the species level. We used mitochondrial cytochrome oxidase I and two nuclear genes, the elongation factor‐1α and LW rhodopsin, to investigate the phylogeny of Sphecidae digger wasps. We focus particularly on the tribe Ammophilini, a clade of nonsocial apoid wasps that exhibit unusually diverse parental care strategies. We analysed a 2232 bp dataset for 40 ammophilines plus nine other taxa from within the remaining Sphecidae. Our Bayesian phylogeny provides strong support for the monophyly of Ammophilini and for the monophyly of all six individual ammophiline genera, except that the position of P. affinis within the genus Podalonia is only weakly supported. The monophyly of some, but not all, previously designated species groups within the genus Ammophila is supported. We discuss the implications of our results for the evolution of morphological traits used previously in ammophiline systematics.  相似文献   

20.
Horismenus parasitoids are an abundant and understudied group of eulophid wasps found mainly in the New World. Recent surveys based on morphological analyses in Costa Rica have quadrupled the number of named taxa, with more than 400 species described so far. This recent revision suggests that there is still a vast number of unknown species to be identified. As Horismenus wasps have been widely described as parasitoids of insect pests associated with crop plants, it is of high importance to properly establish the extant diversity of the genus, in order to provide biological control practitioners with an exhaustive catalog of putative control agents. In this study, we first collected Horismenus wasps from wild Phaseolus bean seeds in Central Mexico and Arizona to assess the genetic relatedness of three morphologically distinct species with overlapping host and geographical ranges. Sequence data from two nuclear and two mitochondrial gene regions uncovered three cryptic species within each of the three focal species (i.e., H. missouriensis, H. depressus and H. butcheri). The monophyly of each cryptic group is statistically supported (except in two of them represented by one single tip in which monophyly cannot be tested). The phylogenetic reconstruction is discussed with respect to differences between gene regions as well as likely reasons for the differences in variability between species.  相似文献   

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