Cladistic analysis of self‐grooming indicates a single origin of eusociality in corbiculate bees (Hymenoptera: Apidae)

Behavioural traits have been used extensively in recent years as an important character source for making phylogenetic inferences. The phylogenetic positions of the members of the Apini subtribe are increasingly being debated, and new characters must be examined. We analysed the presence and absence of certain behavioural patterns, as well as the sequences of some of these patterns, to generate 79 characters. Eleven species comprised the ingroup, and Xylocopini comprised the outgroup. Parsimony analysis showed that the most parsimonious tree was (Euglossina(Bombina(Apina+Meliponina))). This topology is consistent with most studies that use morphological data and the few that use behavioural data, which suggests that advanced eusociality arose only once in a common ancestor of the clade Apina plus Meliponina; however, this hypothesis is inconsistent with our molecular data. Thus we considered behavioural, molecular, and morphological data and recovered the same topology, in which eusociality has a single origin in corbiculate bees.     

Phylogenetic utility of the major opsin in bees (Hymenoptera: Apoidea): a reassessment

Major opsin (LW Rh) DNA sequence has been reported to provide useful data for resolving phylogenetic relationships among tribes of corbiculate bees based on analyses of 502 bp of coding sequence. However, the corbiculate tribes are believed to be of Cretaceous age, and strong support for insect clades of this age from small data sets of nucleotide sequence data has rarely been demonstrated. To more critically assess opsin's phylogenetic utility we generated an expanded LW Rh data set by sequencing the same gene fragment from 52 additional bee species from 24 tribes and all six extant bee families. Analyses of this data set failed to provide substantial support for monophyly of corbiculate bees, for relationships among corbiculate tribes, or for most other well-established higher-level relationships among long-tongued bees. However, monophyly of nearly all genera and tribes is strongly supported, indicating that LW Rh provides useful phylogenetic signal at lower taxonomic levels. When our expanded LW Rh data set is combined with a morphological and behavioral data set for corbiculate bees, the results unambiguously support the traditional phylogeny of the corbiculate bee tribes: (Euglossini + (Bombini + (Meliponini + Apini))). This implies a single origin of advanced eusocial behavior among bees rather than dual origins, as proposed by several recent studies.     

Behavioral Phylogeny of Corbiculate Apidae (Hymenoptera; Apinae), with Special Reference to Social Behavior

The phylogenetic relationships among the four tribes of corbiculate bees (Euglossini, Bombini, Meliponini, and Apini) are controversial. There is substantial incongruence between morphological and molecular data, and the single origin of eusociality is questionable. The use of behavioral characters by previous workers has been restricted to some typological definitions, such as solitary and eusocial. Here, I expand the term "social" to 42 characters and present a tree based only on behavioral characters. The reconstructed relationships were similar to those observed in morphological and "total evidence" analyses, i.e., Euglossini + (Bombini + (Meliponini + Apini)), all of which support a single origin of eusociality.     

Comparative morphology of terminal-instar larvae of Cynipoidea: phylogenetic implications

We describe the external morphology of the terminal-instar larvae of 30 species of Cynipoidea (Hymenoptera), with special reference to the head capsule and mouthparts. Twenty-five of the species belong to the Cynipidae and are gall inducers or phytophagous inquilines (guests) in galls, while five represent different insect-parasitic lineages of the Cynipoidea. Although we find only limited variation in body shape, the head sclerites and mandibles offer many characters of potential phylogenetic value. For instance, the mandibles of the parasitoids have one large pointed tooth, with several smaller dents along the inner margin in core figitids, whereas the phytophagous gall inducers and inquilines have mandibles with two or three blunt teeth of subequal size. The mandibles of inquiline larvae are unique in being covered by vertical striations and in having a dominating, broad second tooth. We summarize the qualitative variation among the studied terminal-instar larvae in terms of 33 morphological characters and one life-history trait and examine the phylogenetic implications of these data by running parsimony analyses under uniform character weights and under implied weights (Goloboff weights). The analysis under uniform weights is poorly resolved but the relationships suggested by the implied-weights analysis are largely congruent with previous analyses of adult morphology and molecular data. The larval data support inclusion of the genus Liposthenes in the Neaylax – Isocolus clade, in agreement with the molecular data but in weak conflict with adult morphology. However, the larval data agree with adult morphology and conflict with the molecular data in supporting monophyly of the inquilines.     

On the Head Morphology and Systematic Position of Helophorus (Coleoptera: Hydrophiloidea: Helophoridae)

External and internal features of the head of adults of Helophorus spp. were examined and described in detail. The 6 species under consideration show very little morphological variation. The only distinctive characters, which characterise groups of species, are the presence or absence of the mandibular retinaculum and the symmetric or asymmetric shape of the ultimate maxillary palpomere. Helophoridae is supported by several autapomorphies, e.g. a group of long hairs on the posterodorsal margin of galeomere II and the distinctly serrate hind margin of the right mola. Characters with a potential phylogenetic relevance are listed, presented as a data matrix and analysed cladistically. The monophyly of Hydrophiloidea + Histeroidea, Hydrophiloidea (excl. Hydraenidae), ((Helophoridae + Hydrochidae) + (Georissidae [+ Epimetopidae?])), Hydrophilidae and Sphaeridiinae was supported in all trees. The position of Spercheus remains ambiguous. It is either the sistergroup of the remaining Hydrophiloidea or of Hydrophilidae. Head structures of adults of Helophoridae and Hydrochidae show a remarkable similarity. The following apomorphic character states are shared by both taxa: dorsal side of labrum divided into 2 areas with different surface structure, mentum with 2 longitudinal ridges. A clade comprising these 2 families + Georissidae (and probably Epimetopidae) is supported by the metallic granulation of the dorsal side of the head capsule and a grooved frontoclypeal suture. The presence of tubular mandibular glands may be a derived groundplan feature of Hydrophiloidea + Histeroidea. The proposed interrelationships are partly in contrast to current hypothesis. The hypothesised character evolution may change, if a more extensive set of taxa (e.g. Horelophinae, Horelophopsinae) and characters, especially larval features are used (e.g. stigmatic atrium). Several derived characteristics of the clades listed above may have been secondarily lost in Hydrophilidae.     

Phylogenetic analysis of the corbiculate Apinae based on morphology of the sting apparatus (Hymenoptera: Apidae)

This study aimed to test the various competing hypotheses regarding the relationships among the four tribes of corbiculate apine bees (Euglossini “orchid bees”, Bombini “bumble bees”, Meliponini “stingless bees”, and Apini “honey bees”) with a completely new set of previously unstudied morphological characters derived from the sting apparatus. The result was one most parsimonious tree of 49 steps, CI = 89, RI = 93 that is perfectly congruent with most studies based on morphological and combined morphological/molecular data, i.e., Euglossini + (Bombini + (Meliponini + Apini)), supporting a well accepted scenario of social evolution for these bees. This data matrix was then combined with other published matrices for this group in order to perform simultaneous analyses. The problem of how to best combine the multiple matrices that did not use the same exemplars was investigated. © The Willi Hennig Society 2007.     

Phylogenetic analysis of four nuclear protein-encoding genes largely corroborates the traditional classification of Bivalvia (Mollusca)

Revived interest in molluscan phylogeny has resulted in a torrent of molecular sequence data from phylogenetic, mitogenomic, and phylogenomic studies. Despite recent progress, basal relationships of the class Bivalvia remain contentious, owing to conflicting morphological and molecular hypotheses. Marked incongruity of phylogenetic signal in datasets heavily represented by nuclear ribosomal genes versus mitochondrial genes has also impeded consensus on the type of molecular data best suited for investigating bivalve relationships. To arbitrate conflicting phylogenetic hypotheses, we evaluated the utility of four nuclear protein-encoding genes-ATP synthase β, elongation factor-1α, myosin heavy chain type II, and RNA polymerase II-for resolving the basal relationships of Bivalvia. We sampled all five major lineages of bivalves (Archiheterodonta, Euheterodonta [including Anomalodesmata], Palaeoheterodonta, Protobranchia, and Pteriomorphia) and inferred relationships using maximum likelihood and Bayesian approaches. To investigate the robustness of the phylogenetic signal embedded in the data, we implemented additional datasets wherein length variability and/or third codon positions were eliminated. Results obtained include (a) the clade (Nuculanida+Opponobranchia), i.e., the traditionally defined Protobranchia; (b) the monophyly of Pteriomorphia; (c) the clade (Archiheterodonta+Palaeoheterodonta); (d) the monophyly of the traditionally defined Euheterodonta (including Anomalodesmata); and (e) the monophyly of Heteroconchia, i.e., (Palaeoheterodonta+Archiheterodonta+Euheterodonta). The stability of the basal tree topology to dataset manipulation is indicative of signal robustness in these four genes. The inferred tree topology corresponds closely to those obtained by datasets dominated by nuclear ribosomal genes (18S rRNA and 28S rRNA), controverting recent taxonomic actions based solely upon mitochondrial gene phylogenies.     

How many marmoset (Primates: Cebidae: Callitrichinae) genera are there? A phylogenetic analysis based on multiple morphological systems

The marmosets, tribe Callitrichini, are the most speciose clade in the subfamily Callitrichinae, containing 21 species. However, there is no consensus among molecular and morphological systematists as to how many genera should be recognized for the group. To test the morphological support for the alternative generic classifications, this study presents a comprehensive phylogenetic analysis. It is the first such analysis to include all 21 species and employ continuous and discrete osteological, pelage and tegument, karyological and vocal characters. This dataset was combined with nucleotide sequences from two mitochondrial and four nuclear regions. Separate analyses showed that, among morphological datasets, osteological characters were best at solving relationships at more inclusive levels, whilst pelage characters were most informative at the interspecific level. This suggests the presence of different transformation rates for the two character sets. When a single most parsimonious tree was obtained using the 83‐character matrix, three main clades were identified, supporting the division of the marmosets into three genera: Callithrix, Cebuella and Mico. The total evidence analysis that included an additional 3481 molecular characters corroborated most of the morphology‐based clades and also supported a three‐genus classification of the marmosets. This is the first morphological study to support an Amazonian marmoset clade (Cebuella + Mico), which is also strongly supported in exclusively molecular phylogenies, and to synonimize Callibella under Mico.     

Phylogenetic synthesis of morphological and molecular data reveals new insights into the higher‐level classification of Tipuloidea (Diptera)

Tipuloidea, the crane flies, are a diverse lineage of true flies (Insecta: Diptera) whose phylogenetic classification and taxonomy remain a challenge. Here we present the results of a quantitative phylogenetic analysis of Tipuloidea based on combined morphological characters (adult, larvae and pupae) and nuclear gene sequence data (28S rDNA and CAD). Forty‐five species, from 44 genera and subgenera, were sampled, representing the four putative families of Tipuloidea (Cylindrotomidae, Limoniidae, Pediciidae and Tipulidae sensu stricto). Analyses of individual datasets, although differing in overall topology, support the monophyly of several major lineages within Tipuloidea. Parsimony and Bayesian analyses using individual morphological and molecular datasets resulted in incongruent topologies. Increased resolution and tree support was obtained when both datasets (morphology and genes) were combined, in both combined evidence parsimony and Bayesian analyses, than when analysed separately. The recovered consensus phylogeny was not consistent with any previously proposed Tipuloidea classification, with previous importance assigned to character states shown here to represent losses and reversals seen as a major factor influencing erroneous classification. The results provided here, together with evidence from previous analyses, were used to append the Tipuloidea classification to supported evolutionary lineages. Tipuloidea is presented as two families: Pediciidae and Tipulidae. Pediciidae is recovered as the sister group to all remaining Tipuloidea. Our current phylogenetic hypothesis is not consistent with the existing subfamilial classification of the ‘Limoniidae’, which is paraphyletic with respect to a well‐supported Tipulinae + Cylindrotominae clade, whereas the three ‘limoniid’ subfamilies are para‐ or polyphyletic. The recognition of ‘Limoniidae’ as a valid monophyletic family is discouraged and the subfamilies of ‘Limoniidae’ are amended and placed within Tipulidae. A revised phylogenetic classification is proposed for the crane flies based on a synthesis of evidence from multiple genes and morphology.     

Phylogeny of Myrmeleontiformia based on larval morphology (Neuropterida: Neuroptera)

The suborder Myrmeleontiformia is a derived lineage of lacewings (Insecta: Neuroptera) including the families Psychopsidae, Nemopteridae, Nymphidae, Ascalaphidae and Myrmeleontidae. In particular, Myrmeleontidae (antlions) are the most diverse neuropteran family, representing a conspicuous component of the insect fauna of xeric environments. We present the first detailed quantitative phylogenetic analysis of Myrmeleontiformia, based on 107 larval morphological and behavioural characters for 36 genera whose larvae are known (including at least one representative of all the subfamilies of the suborder). Four related families were used as outgroups to polarize character states. Phylogenetic analyses were conducted using both parsimony and Bayesian methods. The reconstructions resulting from our analyses corroborate the monophyly of Myrmeleontiformia. Within this clade, Psychopsidae are recovered as the sister family to all the remaining taxa. Nemopteridae (including both subfamilies Nemopterinae and Crocinae) are recovered as monophyletic and sister to the clade comprising Nymphidae + (Myrmeleontidae + Ascalaphidae). Nymphidae consist of two well‐supported clades corresponding to the subfamilies Nymphinae and Myiodactylinae. Our results suggest that Ascalaphidae may not be monophyletic, as they collapse into an unresolved polytomy under the Bayesian analysis. In addition, the recovered phylogenetic relationships diverge from the traditional classification scheme for ascalaphids. Myrmeleontidae are reconstructed as monophyletic, with the subfamilies Stilbopteryginae, Palparinae and Myrmeleontinae. We retrieved a strongly supported clade comprising taxa with a fossorial habit of the preimaginal instars, which represents a major antlion radiation, also including the monophyletic pit‐trap building species.     

A Morphological Basis for Assessing the Phylogeny of the "Tenrecoidea" (Mammalia, Lipotyphla)

A phylogenetic analysis of 35 mammalian taxa focusing on the lipotyphlan family Tenrecidae, based on 193 morphological character states across 71 characters, is undertaken to test several hypotheses of lipotyphlan relationships, including monophyly of the Lipotyphla, Tenrecidae, Malagasy Tenrecidae, and Caribbean Lipotyphla. Explicit tests of these hypotheses are central to understanding larger issues concerning Malagasy and Caribbean biogeography, in addition to mammalian phylogeny. Methodologically, several different parameters are created with which to determine the sensitivity of resulting clades to initial assumptions about a posteriori character weighting, missing data, and multistate character ordering. Clades produced by this data set that appear despite perturbations in these parameters and that are supported by other confidence-assessment techniques contradict Malagasy tenrecid monophyly and the association of soricids with the Caribbean taxon Solenodon. Results regarding tenrecid and lipotyphlan monophyly are more ambiguous and depend on certain assumptions regarding treatment of character ordering, weighting, and missing data. An alternative phylogeny supporting an African mammal clade receives no support from the data set discussed herein.     

Molecular tests of phylogenetic taxonomies: a general procedure and example using four subfamilies of the lizard family Iguanidae

A general procedure is described for examining when results of molecular phylogenetic analyses warrant formal revision of taxonomies constructed using morphological characters. We illustrate this procedure with tests of monophyly for four subfamilies in the lizard family Iguanidae using 1561 aligned base positions (838 phylogenetically informative) of mitochondrial DNA sequences, representing coding regions for eight tRNAs, ND2, and portions of ND1 and COI. Ten new sequences ranging in length from 1732 to 1751 bases are compared with 12 previously reported sequences and 67 morphological characters (54 phylogenetically informative) from the literature. New morphological character states are provided for Sator. Phylogenies derived from the molecular and combined data are in agreement but both conflict with phylogenetic inferences from the morphological data alone. Strong support is found for the monophyly of the subfamilies Crotaphytinae and Phrynosomatinae. Monophyly of the Iguaninae is weakly supported in each analysis. All analyses suggest that the Tropidurinae is not monophyletic but the hypothesis of monophyly cannot be rejected. A phylogenetic taxonomy is proposed in which the Tropidurinae* is maintained as a metataxon (denoted with an asterisk), for which monophyly has not been demonstrated. Within the Phrynosomatinae, the close relationship of Sator and Sceloporus is questioned and an alternative hypothesis in which Sator is the sister taxon to a clade comprising Petrosaurus, Sceloporus, and Urosaurus is presented. Statistical tests of monophyly provide a powerful way to evaluate support for taxonomic groupings. Use of the metataxon prevents premature taxonomic rearrangements where support is lacking.     

The osteology and phylogeny of the Hawaiian finch radiation (Fringillidae: Drepanidini), including extinct taxa

The monophyly and phylogeny of the adaptive radiation of Hawaiian finches (Fringillidae: Drepanidini; honeycreepers, auct.) were studied using parsimony analysis of comparative osteology, combined with Templeton (Wilcoxon signed‐ranks) tests of alternative phylogenetic hypotheses. Eighty‐four osteological characters were scored in 59 terminal taxa of drepanidines, including 24 fossil forms, and in 30 outgroup species. The optimal phylogenetic trees show considerable agreement, and some conflict, with independently derived ideas about drepanidine evolution. The monophyly of a large Hawaiian radiation was upheld, although one fossil taxon from Maui fell outside the drepanidine clade. The finch‐billed species were placed as basal drepanidine taxa, and continental cardueline finches (Carduelini) were identified as the radiation's closest outgroups. The study found anatomical as well as phylogenetic evidence that the radiation had a finch‐billed ancestor. The optimal trees identify the red‐and‐black plumage group as a clade, and suggest that the tubular tongue evolved only once in the radiation. Because comparative osteology provides too few characters to strongly support all the nodes of the tree, it was helpful to evaluate statistical support for alternative hypotheses about drepanidine relationships using the Templeton test. Among the alternatives that received significant statistical support are a relationship of the drepanidines with cardueline finches rather than with the Neotropical honeycreepers (Thraupini), classification of the controversial genera Paroreomyza and Melamprosops as drepanidines, and a secondary loss of the tubular tongue in Loxops mana. The hypothesis of monophyly for all the Hawaiian taxa in the study was not rejected statistically. The study provides a framework for incorporating morphological and palaeontological information in evolutionary studies of the Drepanidini. © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 141 , 207–255.     

Colony size evolution and the origin of eusociality in corbiculate bees (Hymenoptera: Apinae)

Recently, it has been proposed that the one of the main determinants of complex societies in Hymenoptera is colony size, since the existence of large colonies reduces the direct reproductive success of an average individual, given a decreased chance of being part of the reproductive caste. In this study, we evaluate colony size evolution in corbiculate bees and their relationship with the sociality level shown by these bees. Specifically i) the correlation between colony size and level of sociality considering the phylogenetic relationship to evaluate a general evolutionary tendency, and ii) the hypothetical ancestral forms of several clades within a phylogeny of corbiculate bees, to address idiosyncratic process occurring at important nodes. We found that the level of social complexity in corbiculate bees is phylogenetically correlated with colony size. Additionally, another process is invoked to propose why colony size evolved concurrently with the level of social complexity. The study of this trait improves the understanding of the evolutionary transition from simple to complex societies, and highlights the importance of explicit probabilistic models to test the evolution of other important characters involved in the origin of eusociality.     

Higher level phylogeny for the passion-vine butterflies (Nymphalidae, Heliconiinae) based on early stage and adult morphology

A higher level phylogeny for the passion-vine butterflies (Nymphalidae, Heliconiinae) was generated by cladistic analysis of 146 morphological characters from all life stages. The 24 species studied were selected representatives of the ten currently accepted genera of the sub-tribe Heliconiiti. Analyses of only characters from larvae and pupae did not produce well resolved trees. However, some characters of the immature stages provided critical support for the monophyly of two clades. Analysis of only adult characters yielded a tree that closely resembled that obtained from all data combined. The phylogeny here derived from the combined analysis of early stage and adult characters differed in topology from all previously proposed hypotheses, and supported the monophyly of all currently recognized genera. Characters supporting each clade are described and illustrated, and various hypotheses of phylogenetic relatedness of passion-vine butterfly taxa are discussed.     

Evolution and phylogeny of the scarab subtribe Anisopliina (Coleoptera: Scarabaeidae: Rutelinae: Anomalini)

Abstract.  The subtribe Anisopliina (Scarabaeidae: Rutelinae: Anomalini) is associated with grasses, and its species are distributed in the Palaearctic, Oriental, Ethiopian, Nearctic and Neotropical biogeographical regions. Phylogenetic analysis of adult morphological characters was conducted to examine the monophyly and classification of the group, as well as to examine characters associated with grass pollinivory and graminivory. We review the biology, phylogeny and classification of the Anisopliina and provide an overview of each genus. The analysis of ninety-one morphological characters using parsimony does not support the monophyly of the subtribe Anisopliina. Instead, the results provide support for a group referred to here as the anisopliine clade, a circum-Mediterranean group, forming an internal clade within the well-supported tribe Anomalini. Sister group relationships are discussed, possibly being associated with a New World anomaline taxon. Character states associated with grass herbivory, including mouthpart and leg characters, are discussed based on the phylogenetic analysis. Within the Anomalini, an evolutionary shift from generalized leaf feeding to grass associations and grass pollen feeding is supported.     

Phylogenetics, biogeography and classification of, and character evolution in, gamebirds (Aves: Galliformes): effects of character exclusion, data partitioning and missing data

The phylogenetic relationships, biogeography and classification of, and morpho‐behavioral (M/B) evolution in, gamebirds (Aves: Galliformes) are investigated. In‐group taxa (rooted on representatives of the Anseriformes) include 158 species representing all suprageneric galliform taxa and 65 genera. The characters include 102 M/B attributes and 4452 nucleic acid base pairs from mitochondrial cytochrome b (CYT B), NADH dehydrogenase subunit 2 (ND2), 12S ribosomal DNA (12S) and control region (CR), and nuclear ovomucoid intron G (OVO‐G). Analysis of the combined character data set yielded a single, completely resolved cladogram that had the highest levels of jackknife support, which suggests a need for a revised classification for the phasianine galliforms. Adding 102 M/B characters to the combined CYT B and ND2 partitions (2184 characters) decisively overturns the topology suggested by analysis of the two mtDNA partitions alone, refuting the view that M/B characters should be excluded from phylogenetic analyses because of their relatively small number and putative character state ambiguity. Exclusion of the OVO‐G partition (with > 70% missing data) from the combined data set had no effect on cladistic structure, but slightly lowered jackknife support at several nodes. Exclusion of third positions of codons in an analysis of a CYT B + ND2 partition resulted in a massive loss of resolution and support, and even failed to recover the monophyly of the Galliformes with jackknife support. A combined analysis of putatively less informative, “non‐coding” characters (CYT B/ND2 third position sites + CR +12S + OVO‐G sequences) yielded a highly resolved consensus cladogram congruent with the combined‐evidence cladogram. Traditionally recognized suprageneric galliform taxa emerging in the combined cladogram are: the families Megapodiidae (megapodes), Cracidae (cracids), Numididae (guineafowls), Odontophoridae (New World quails) and Phasianidae (pheasants, pavonines, partridges, quails, francolins, spurfowls and grouse) and the subfamilies Cracinae (curassows, chachalacas and the horned guan), Penelopinae (remaining guans), Pavoninae sensu lato (peafowls, peacock pheasants and argus pheasants), Tetraoninae (grouse) and Phasianinae (pheasants minus Gallus). The monophyly of some traditional groupings (e.g., the perdicinae: partridges/quails/francolins) is rejected decisively, contrasted by the emergence of other unexpected groupings. The most remarkable phylogenetic results are the placement of endemic African galliforms as sisters to geographically far‐distant taxa in Asia and the Americas. Biogeographically, the combined‐data cladogram supports the hypothesis that basal lineages of galliforms diverged prior to the Cretaceous/Tertiary (K‐T) Event and that the subsequent cladogenesis was influenced by the break‐up of Gondwana. The evolution of gamebirds in Africa, Asia and the Americas has a far more complicated historical biogeography than suggested to date. With regard to character evolution: spurs appear to have evolved at least twice within the Galliformes; a relatively large number of tail feathers (≥ 14) at least three times; polygyny at least twice; and sexual dimorphism many times. © The Willi Hennig Society 2006.     

白鱼属鱼类的系统发育(鲤形目:鲤科)

白鱼属(Anabarilius Cockerell)是我国特有的鲤科鱼类,主要分布于云南东部、中部和四川南部。分布范围不大,但种的分化强烈。 Regan(1904,1908, 1918)最先报道四种白鱼,但均置于Barilius属。尔后,Cockerell(1923)以Barilius andersoni Regan为模式种建立白鱼属;Nichols(1927)、张孝威(1944)、易伯鲁、吴清江(1964)、刘振华、何纪昌(1983)先后记述了部分种和亚种。陈银瑞、褚新洛(1980)在系统整理的基础上,澄清了白鱼属属级分类上的混乱,记述了三新种和一新亚种。何纪昌、王重光(1984)用数值分类法探讨了白鱼属的分类,报道了二新种和一新亚种。至此,白鱼属共包括15个种和亚种。     

A comprehensive phylogeny of the bumble bees (Bombus)

Bumble bees ( Bombus Latreille) occupy a wide diversity of habitats, from alpine meadows to lowland tropical forest, yet they appear to be similar in morphology throughout their range, suggesting that behavioural adaptations play a more important role in colonizing diverse habitats. Notwithstanding their structural homogeneity, bumble bees exhibit striking inter- and intraspecific variation in colour pattern, purportedly the outcome of mimetic evolution. A robust phylogeny of Bombus would provide the framework for elucidating the history of their wide biogeographical distribution and the evolution of behavioural and morphological adaptations, including colour pattern. However, morphological studies of bumble bees have discovered too few phylogenetically informative characters to reconstruct a robust phylogeny. Using DNA sequence data, we report the first nearly complete species phylogeny of bumble bees, including most of the 250 known species from the 38 currently recognized subgenera. Bayesian analysis of nuclear (opsin, EF-1α, arginine kinase, PEPCK) and mitochondrial (16S) sequences results in a highly resolved and strongly supported phylogeny from base to tips, with clear-cut support for monophyly of most of the conventional morphology-based subgenera. Most subgenera fall into two distinct clades ( short-faced and long-faced ) associated broadly with differences in head morphology. Within the short-faced clade is a diverse New World clade, which includes nearly one-quarter of the currently recognized subgenera, many of which are restricted to higher elevations of Central and South America. The comprehensive phylogeny provides a firm foundation for reclassification and for evaluating character evolution in the bumble bees.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 161–188.