寄生蜂核型特征及其在分类上的应用

回顾了寄生性膜翅目昆虫（寄生蜂）染色体数目和核型的研究情况及其在寄生蜂分类中的应用,并对寄生蜂核型分类的前景作了讨论。核型对众多的寄生性膜翅目类群来说可以提供许多分类上的信息,并在种类研究上作用最为显著。通过核型数据的进一步积累和染色体显带方法的进一步应用,寄生蜂的核型分类学将得到进一步的发展。     

Karyotypes of parasitic Hymenoptera: Diversity, evolution and taxonomic significance

Haploid chromosome numbers (n) of parasitic Hymenoptera (= traditional Parasitica + Chrysidoidea) vary from 2 to 23. However, this range can be subdivided into three intervals with n= 14–23 (less derived parasitic wasps, e.g., some Ichneumonidae and Braconidae as well as Gasteruptiidae), 8–13 (many other parasitic Hymenoptera) and 2–7 (Dryinidae, the majority of Chalcidoidea and some advanced Braconidae, e.g. Aphidiinae). The symmetric karyotype with a relatively high chromosome number (n= 14–17) and the prevalence of biarmed chromosomes must be considered as a groundplan feature of parasitic Hymenoptera. Independent reductions of chromosome numbers (n≤ 10–11) occurred in some groups of the superfamily Ichneumonoidea as well as in the common ancestor of the Proctotrupoidea sensu lato, Ceraphronoidea, Cynipoidea and Chalcidoidea. Further multiple decreases in chromosome numbers (n≤ 4–6) took place in some Braconidae, various lineages of the superfamily Chalcidoidea as well as in the family Dryinidae. Two main trends prevailed in the karyotype evolution of parasitic wasps: the reduction of chromosome numbers (mainly due to tandem fusions and less frequently due to centric ones) and karyotypic dissymmetrization (through an increase in size differentiation of chromosomes and/or in the share of acrocentrics in a chromosome set). Although karyotypic features of parasitic Hymenoptera can be used for solving taxonomic problems at various levels, this method is the most effective at the species level.     

Chromosomal evolution in the lizard genus Varanus (reptilia).

The karyotypes have been determined of 16 of the 32 species of the genus Varanus, including animals from Africa, Israel, Malaya and Australia. A constant chromosome number of 2n = 40 was observed. The karyotype is divided into eight pairs of large chromosomes and 12 paris of microchromosomes. A series of chromosomal rearrangements have become established in both size groups of the karyotype and are restricted to centromers shifts, probably caused by pericentric inversion. Species could be placed in one of six distinct karyotype groups which are differentiated by these rearrangements and whose grouping does not always correspond with the current taxonomy. An unusual sex chromosome system of the ZZ/ZW type was present in a number of the species examined. The evolutionary significance of these chromosomal rearrangements, their origin and their mode of establishment are discussed and related to the current taxonomic groupings. The most likely phylogenetic model based on chromosome morphology, fossil evidence and the current distribution of the genus Varanus is presented.     

Evolutionary trends in the chromosome numbers of swarm-founding social wasps

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.     

Low courtship song variation in south and southeastern Brazilian populations of Drosophila meridionalis (Diptera, Drosophilidae)

Drosophila meridionalis is a cactus-breeding species with a wide distribution in South America. Most populations of this species are geographically isolated, what provides a promising scenario for studying evolution. Former studies of this species revealed a remarkable karyotypic variation among its populations. Up to six distinct metaphase chromosomes were described, showing that this species is polymorphic at least at the chromosomal level. In order to elucidate the taxonomic status of populations showing different metaphase chromosomes, we analyzed the courtship song of five populations of D. meridionalis in South and Southeastern Brazil. In addition, we analyzed the metaphase chromosomes of each population. Our results show that, despite the two karyotype observed, most courtship song parameters did not vary among the populations. Altogether, our results suggest that D. meridionalis from South and Southeastern Brazil represents one species with an inter-population chromosomal variability.     

Chromosomal apparatus of trematodes of the general Diplostomum and Tylodelphys (Strigeidida, Diplostomatidae) and its taxonomic significance

The chromosomal apparatus of three species of trematodes of Diplostomum (D. spathaceum, D. indistinctum, D. mergi) and one species Tylodelphys (T. clavata) is described. It has been suggested that karyological investigations can be used for taxonomic purposes. The opinion has been expressed that the number of chromosomes in the karyotype of these trematodes plays no taxonomic role. Of great taxonomic significance are the size and morphology of chromosomes. Differences in karyotypes of trematodes belonging to different species and genera were revealed by these characters. The absence of close dependence between morphology of adult and specific peculiarities of karyotypes in species of the same genus enables in some cases the use of chromosomal apparatus as a most reliable criterion for a differentiation of close species.     

New insights into and limitations of the molecular phylogeny in the taxon-rich land snail genus Montenegrina (Mollusca: Gastropoda: Clausiliidae)

Rock-dwelling gastropods are usually patchily distributed in limestone habitats, presumably have low active and passive dispersal ability and often represent narrow-ranged endemic taxa. Their current taxonomy is predominantly shell morphology based, and it remains unknown whether the morphologically differentiated and geographically separated populations represent phylogenetic clades. In this study, we analysed the hyperdiverse, terrestrial door snail genus Montenegrina. Based on the current taxonomy defined by shell morphology, it contains 29 species and 106 subspecies distributed in the Balkan region. The constructed phylogenetic tree using three mitochondrial markers was used to test whether it agrees with the current taxonomy. In this comprehensive tree, about half of the species and subspecies are monophyletic. Some of the paraphylies could be reasonably resolved by taxonomic changes; that is, some subspecies should be reassigned or raised to species level. Other incongruencies probably arose due to introgression even between distant clades. The histone genes turned out to be unsuitable for elucidating the phylogeny of Montenegrina. In the species-delimitation tests, considerably more molecular operational taxonomic units were delimited than the number of presently described species. The present data indicate that (a) shell morphology-based taxonomy and taxon recognition can be problematic in such a large and morphologically highly variable genus; (b) the potential error due to incomplete sampling presents a problem in a genus as variable as Montenegrina; (c) multi-locus analyses should be conducted to arrive at a better basis for species delimitation; and (d) integrative approaches including genetic as well as morphological/anatomical data from a comprehensive geographic sample are necessary.     

Skeletal morphology of Opius dissitus and Biosteres carbonarius (Hymenoptera: Braconidae), with a discussion of terminology

The Braconidae, a family of parasitic wasps, constitute a major taxonomic challenge with an estimated diversity of 40,000 to 120,000 species worldwide, only 18,000 of which have been described to date. The skeletal morphology of braconids is still not adequately understood and the terminology is partly idiosyncratic, despite the fact that anatomical features form the basis for most taxonomic work on the group. To help address this problem, we describe the external skeletal morphology of Opius dissitus Muesebeck 1963 and Biosteres carbonarius Nees 1834, two diverse representatives of one of the least known and most diverse braconid subfamilies, the Opiinae. We review the terminology used to describe skeletal features in the Ichneumonoidea in general and the Opiinae in particular, and identify a list of recommend terms, which are linked to the online Hymenoptera Anatomy Ontology. The morphology of the studied species is illustrated with SEM-micrographs, photos and line drawings. Based on the examined species, we discuss intraspecific and interspecific morphological variation in the Opiinae and point out character complexes that merit further study.     

塞威氏苹果Malus sieversii（Ldb.）Roem花粉形态的研究

对采自75个不同居群的塞威氏苹果（Malus sieversii (Ldb.)Roem）的花粉形态进行了观察。结果表明：塞威氏苹果花粉萌发孔为三孔沟类型,外壁具条纹状纹饰,条纹间常具小穴或穿孔。根据外壁纹饰的差异将塞威氏苹果花粉划分为六个类型。同时根据花粉资料,探讨了塞威氏苹果的某些分类学和系统学问题。     

Species limits in butterflies (Lepidoptera: Nymphalidae): reconciling classical taxonomy with the multispecies coalescent

Species delimitation is at the core of biological sciences. During the last decade, molecular‐based approaches have advanced the field by providing additional sources of evidence to classical, morphology‐based taxonomy. However, taxonomy has not yet fully embraced molecular species delimitation beyond threshold‐based, single‐gene approaches, and taxonomic knowledge is not commonly integrated into multilocus species delimitation models. Here we aim to bridge empirical data (taxonomic and genetic) with recently developed coalescent‐based species delimitation approaches. We use the multispecies coalescent model as implemented in two Bayesian methods (dissect/stacey and bp&p ) to infer species hypotheses. In both cases, we account for phylogenetic uncertainty (by not using any guide tree) and taxonomic uncertainty (by measuring the impact of using a priori taxonomic assignments to specimens). We focus on an entire Neotropical tribe of butterflies, the Haeterini (Nymphalidae: Satyrinae). We contrast divergent taxonomic opinion – splitting, lumping and misclassifying species – in the light of different phenotypic classifications proposed to date. Our results provide a solid background for the recognition of 22 species. The synergistic approach presented here overcomes limitations in both traditional taxonomy (e.g. by recognizing cryptic species) and molecular‐based methods (e.g. by recognizing structured populations, and not raising them to species). Our framework provides a step forward towards standardization and increasing reproducibility of species delimitations.     

我国直翅目昆虫细胞分类学研究现状

从研究材料、研究方法、染色体研究在昆虫分类上的应用 3个方面对我国昆虫尤其是直翅目昆虫的细胞分类学研究情况作了简要的概述 ,并着重论述了直翅目昆虫的核型及带型的研究现状 ,同时对我国直翅目昆虫分类学的发展前景进行了展望     

Review: Taxonomy of Nitella (Charales, Charophyceae) based on comparative morphology of oospores and multiple DNA marker phylogeny using cultured material

The genus Nitella is the largest group in the Charales and has the highest diversity of vegetative and oospore morphologies. In his worldwide monograph on the Charales, R. D. Wood characterized the sections and species of Nitella mainly on the basis of vegetative morphology and treated oospore wall morphology as diagnostic at the infraspecific level. Therefore, many species of Nitella exhibiting distinct external morphology of the oospore wall (EMOW) were reduced to infraspecific rank and only 53 of 204 species previously described were recognized within Nitella . However, recent morphological and molecular phylogenetic studies have demonstrated the phylogenetic validity of using EMOW for diagnosing some species of Nitella . More recently, a scanning electron microscopy (SEM) study of internal morphology of the oospore wall (IMOW) and multiple DNA marker analyses using both nuclear and chloroplast gene sequences were conducted to improve our understanding of the taxonomy of Nitella at the species level, on the basis of cultured material of a large number of species. Multiple DNA marker analyses resolved detailed and robust phylogenetic relationships within the genus and demonstrated the taxonomic and phylogenetic significance of IMOW. In addition, they supported the taxonomic decisions based on differences in oospore morphology, suggesting that an integrated approach, involving both SEM studies of the EMOW and IMOW and multiple DNA marker analyses, is appropriate to address problems at lower taxonomic levels within the genus, as well as to construct a natural taxonomic system for the genus. In this paper, recent morphological and molecular phylogenetic studies are reviewed and recent improvements in taxonomy of Nitella are summarized. Moreover, the evolution of morphological features and phylogenetic relationships within Nitella are discussed, focusing especially on oospore morphology.     

中国西南地区鹿药属4种15居群核型研究

对产于中国西南部的鹿药属（Maianthemum）4种植物进行了细胞学研究,包括染色体数目,多倍化,非整倍性和随体染色体,以及核型不对称性和核型进化。结果表明：1）除了在云南丽江采集的Maianthemum tatsienensis染色体数目为2n=72之外,其余的居群全为2n=36;2）核型在居群间存在变异,特别是在具中部染色体和近中部染色体的数目以及随体染色体的数目和位置上。此外,M．nanchuanense和M．szechuanicum的核型是首次报道,B染色体也是首次在该属中发现。我们推测鹿药属的进化方式包括频繁的染色体畸变以及不同水平上的多倍化,而中国西南部是该属的分化中心。     

Comparative morphological analysis of compound eye miniaturization in minute hymenoptera

Due to their small size, diminutive parasitic wasps are outstanding subjects for investigating aspects of body miniaturization. Information on minute compound eyes is still scarce, and we therefore investigated eye morphology in one of the smallest known hymenopteran species Megaphragma mymaripenne (body size 0.2 mm) relative to Anaphes flavipes (body size 0.45 mm) and compared the data with available information for Trichogramma evanescens (body size 0.4 mm). The eyes of all three species are of the apposition kind, and each ommatidium possesses the typical cellular organization of ommatidia found in larger hymenopterans. Compound eye miniaturization does not therefore involve a reduction in cell numbers or elimination of cell types. Six size-related adaptations were detected in the smallest eyes investigated, namely a) a decrease in the radius of curvature of the cornea compared with larger hymenopterans; b) the lack of extensions to the basal matrix from secondary pigment cells; c) the interlocking arrangement of the retinula cell nuclei in neighboring ommatidia; d) the distal positions of retinula cell nuclei in M. mymaripenne; e) the elongated shape of retinula cell pigment granules of both studied species; and f) an increase in rhabdom diameter in M. mymaripenne compared with A. flavipes and T. evanescens. The adaptations are discussed with respect to compound eye miniaturizations as well as their functional consequences based on optical calculations.     

DNA evidence shows vocalizations to be a better indicator of taxonomic limits than plumage patterns in Zimmerius tyrant-flycatchers

We investigated mitochondrial and nuclear DNA in the small tyrant-flycatcher genus Zimmerius (Tyrannidae) and show that molecular data are in strong disagreement with morphology-based taxonomy, but in good concordance with vocal characters. Our molecular data identified two independent cases of well-supported polyphyletic species arrangements within this genus that indicate the following taxonomic changes: elevation of Z. acer and Z. albigularis to species level, separation of northern populations of Z. chrysops as a species and inclusion of southern populations of Z. chrysops into Z. viridiflavus. Although polyphyly has rarely been encountered in bird systematics it has previously been shown for two other tyrannid genera and suggests that tyrannid taxonomy may be poorly resolved, presumably as a consequence of the conserved plumage patterns observed in many tyrannid genera. Our study suggests that vocalizations can be a better indicator of taxonomic limits than plumage pattern in tyrannids.     

Uniformity of karyotypes in Ligularia (Asteraceae: Senecioneae), a highly diversified genus of the eastern Qinghai–Tibet Plateau highlands and adjacent areas

Ligularia , a highly diversified genus in the eastern Qinghai–Tibet Plateau and adjacent areas, was chosen as a suitable subject in which to study speciation patterns in this 'hot spot' area at the chromosomal level. Chromosome numbers and karyotypes were studied in 23 populations of 14 species, most of which are endemic to this area. The basic number x  = 29 was confirmed for all species. Ligularia virgaurea was found to have diploid and triploid cytotypes, 2 n  = 58 and 87. Other species are only diploid, with 2 n  = 58. The karyotypes of all populations within any species, and all species spanning most sections and covering most of the morphological range in Ligularia , are very similar to each other, belonging to type 2A according to Stebbin's classification. This karyotype was also found in its close allies, e.g. Cremanthodium , Ligulariopsis , Parasenecio , and Sinacalia . Aneuploid reduction of chromosome number from 2 n  = 60 to 58 and karyotypic variation was found in Ligularia and its allies. Such a chromosomal pattern with few polyploids infers that variation of karyotype structure at the diploid level seems to be the predominant feature of chromosomal evolution in this group and sympatric speciation via hybridization and polyploidization has played a minor role in its species diversity.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 144 , 329–342.     

DNA barcodes as a tool in biodiversity research: testing pre-existing taxonomic hypotheses in Delphic Apollo butterflies (Lepidoptera,Papilionidae)

Numerous studies have demonstrated that DNA barcoding is an effective tool for detecting DNA clusters, which can be viewed as operational taxonomic units (OTUs), useful for biodiversity research. Frequently, the OTUs in these studies remained unnamed, not connected with pre-existing taxonomic hypotheses, and thus did not really contribute to feasible estimation of species number and adjustment of species boundaries. For the majority of organisms, taxonomy is very complicated with numerous, often contradictory interpretations of the same characters, which may result in several competing checklists using different specific and subspecific names to describe the same sets of populations. The highly species-rich genus Parnassius (Lepidoptera: Papilionidae) is but one example, such as several mutually exclusive taxonomic systems have been suggested to describe the phenotypic diversity found among its populations. Here we provide an explicit flow chart describing how the DNA barcodes can be combined with the existing knowledge of morphology-based taxonomy and geography (sympatry versus allopatry) of the studied populations in order to support, reject or modify the pre-existing taxonomic hypotheses. We then apply this flow chart to reorganize the taxa within the Parnassius delphius species group, solving long-standing taxonomic problems.     

Species coherence in the face of karyotype diversification in holocentric organisms: the case of a cytogenetically variable sedge (Carex scoparia,Cyperaceae)

Background and Aims

The sedge genus Carex, the most diversified angiosperm genus of the northern temperate zone, is renowned for its holocentric chromosomes and karyotype variability. The genus exhibits high variation in chromosome numbers both among and within species. Despite the possibility that this chromosome evolution may play a role in the high species diversity of Carex, population-level patterns of molecular and cytogenetic differentiation in the genus have not been extensively studied.

Methods

Microsatellite variation (11 loci, 461 individuals) and chromosomal diversity (82 individuals) were investigated in 22 Midwestern populations of the North American sedge Carex scoparia and two Northeastern populations.

Key Results

Among Midwestern populations, geographic distance is the most important predictor of genetic differentiation. Within populations, inbreeding is high and chromosome variation explains a significant component of genetic differentiation. Infrequent dispersal among populations separated by >100 km explains an important component of molecular genetic and cytogenetic diversity within populations. However, karyotype variation and correlation between genetic and chromosomal variation persist within populations even when putative migrants based on genetic data are excluded.

Conclusions

These findings demonstrate dispersal and genetic connectivity among widespread populations that differ in chromosome numbers, explaining the phenomenon of genetic coherence in this karyotypically diverse sedge species. More generally, the study suggests that traditional sedge taxonomic boundaries demarcate good species even when those species encompass a high range of chromosomal diversity. This finding is important evidence as we work to document the limits and drivers of biodiversity in one of the world''s largest angiosperm genera.     

Oral Fimbriae and Papillae in Parasitic Lampreys (Petromyzontiformes)

There are 38 species of living lampreys, 18 parasitic and 20 nonparasitic. The parasitic species feed as adults, while the nonparasitic do not. The taxonomy and systematics of the living lampreys is based primarily on dentition characters. Since the number of oral fimbriae and oral papillae have never been systematically investigated in lampreys, we compared them in 17 of the 18 parasitic lampreys to assess their usefulness as taxonomic characters. Both showed little variation with total length and sex within a species, while exhibiting greater variation between species. Parasitic species belonging to the three lamprey families could be distinguished using the number of oral fimbriae: southern hemisphere Mordaciidae (0) and Geotriidae (55–65) and northern hemisphere Petromyzontidae (81–144). However, the taxonomic usefulness of the two characters at the species level was limited. Ten out of the 17 species of parasitic lampreys were placed into four distinct groups based on their numbers of oral fimbriae, and only in two of these was the character diagnostic. Twelve out of the 17 species were placed into two distinct groups based on their numbers of oral papillae and for none of these was the character diagnostic. Blood feeders and intermediate feeders (blood + flesh) were shown to have significantly higher numbers of oral fimbriae than flesh feeders. The higher numbers of oral fimbriae in the former two types of feeders were presumed to be linked to their greater need to create a good seal for feeding purposes. Blood feeders were also shown to have significantly higher numbers of oral papillae than either intermediates or flesh feeders. The higher numbers of oral papillae in the blood feeders were presumed to be linked to their greater need to find suitable attachment sites for feeding purposes.