EVOLUTION OF DIFFERENT GALL TYPES IN WILLOW-FEEDING SAWFLIES (HYMENOPTERA: TENTHREDINIDAE)

The sawflies that feed on the plant family Salicaceae can be divided into eight informal groups based on larval feeding habit or gall type: (1) species with free-living larvae; (2) leaf folders; (3) leaf blade gallers; (4) apical leaf gallers; (5) basal leaf gallers; (6) midrib and petiole gallers; (7) stem gallers; and (8) bud gallers. It has been proposed that the galling habit evolved from free-living larvae via leaf folders, and that the different gall types evolved gradually in the sequence mentioned above. Thus, the galling site would have “wandered” from the leaf margin toward the stem as a result of gradual changes in oviposition site preference. Allozyme data from eight informative loci were used to reconstruct the phylogeny of 18 representative sawfly species. The results suggest that indeed leaf folders seem to be a basal group; leaf blade gallers evolved independently of the other true gallers; apical and basal leaf gallers are not the ancestors of petiole and bud gallers, but they may share a common galling ancestor; bud gallers evolved from midrib/petiole gallers; and stem gallers are polyphyletic. The cause for the observed wandering of the galling site could be intraspecific competition due to a possible “nutrient shading effect” of galls situated closer to the host plant's main vascular system.     

COMPARATIVE ANALYSIS OF GALL MORPHOLOGY IN AUSTRALIAN GALL THRIPS: THE EVOLUTION OF EXTENDED PHENOTYPES

We used a combination of morphometric, phylogenetic, and life-history information to analyze the evolution and possible adaptive significance of gall morphology in a clade of 24 species of gall-inducing thrips (Insecta: Thysanoptera) on Australian Acacia trees. Principal components analysis revealed that galls varied in morphology along two main axes, spherical versus elongate (PC1) and general size (PC2). A high degree of conservation of gall shape on an independently derived phylogeny of the insects and the presence of nine species of Acacia each bearing two or three morphologically disparate gall forms induced by different thrips species indicate that interspecific variation in gall form is determined predominantly by the insects. Character optimization of PC1 on the phylogeny of gall thrips suggested that the ancestral gall form was a simple roll or curl. The diversification of gall form involved four main processes: (1) the convergent evolution of relatively spherical galls in two clades; (2) the evolution of small elongate and hemispherical galls in one clade; (3) the evolution of a lobed interior in a species with a spherical gall and multiple within-gall generations; and (4) the evolution of intraspecific gall polymorphism in a clade of apparent sibling species. Comparative analyses indicated that gall sphericity was associated with the presence of physogastry (foundress hyperfecundity) and that small elongate and hemispherical forms may be associated with the presence of multiple generations in a gall and, perhaps, with the presence of soldier castes. The evolution of a lobed interior in one species, which greatly increases inner surface area, coincided with the evolution of multiple generations. In the clade with intraspecific gall polymorphism in some species, patterns of intraspecific variation mirror patterns of interspecific variation within the clade as a whole. This is the first study to analyze the evolution of gall size and shape in a phylogenetic context and to investigate the life-history correlates of evolutionary changes in gall form. Taken together, our findings indicate that the main selective pressures driving the evolution of gall form in Australian gall thrips on Acacia involve inner surface area to volume relationships, which change in concert with foundress fecundity and the number of within-gall generations.     

SELFISH LARVAE: DEVELOPMENT AND THE EVOLUTION OF PARASITIC BEHAVIOR IN THE HYMENOPTERA

Queens of hymenopteran social parasites manipulate the workers of other social species into raising their offspring. However, nonconspecific brood care may also allow the parasite larvae to control their own development to a greater extent than possible in nonparasitic species. An evolutionary consequence of this may be the loss of the parasite's worker caste if the larvae can increase their fitness by developing into sexuals rather than workers. We argue that this loss is particularly likely in species in which there is little inclusive fitness benefit in working. Retention of a worker caste correlates with characteristics that increase the fitness of working relative to becoming a sexual, such as worker-production of males, high intracolony relatedness, and seasonal environments where the hosts of potential parasite queens are not always available. Further evidence strongly suggests that when the worker caste is evolutionarily lost in perennial species like ants, it disappears rapidly and through a reduction in caste threshold and queen size, so that parasite larvae become queens with less food than required to produce host workers. This evolutionary process, however, appears to lower overall population fitness, resulting in workerless parasite species having small populations and being geographically restricted. Conversely, in annual species like bees and wasps, workerless social parasitism evolves with no size reduction in queens, which is consistent with an expected lower level of queen/offspring conflict.     

PHYLOGENETIC ANALYSIS OF TRAIT EVOLUTION AND SPECIES DIVERSITY VARIATION AMONG ANGIOSPERM FAMILIES

Angiosperm families differ greatly from one another in species richness (S). Previous studies have attributed significant components of this variation to the influence of pollination mode (biotic/abiotic) and growth form (herbaceous/woody) on speciation rate, but these results suffer difficulties of interpretation because all the studies ignored the phylogenetic relationships among families. We use a molecular phylogeny of the angiosperm families to reanalyse correlations between S and family-level traits and use reconstructions of trait evolution to interpret the results. We confirm that pollination mode and growth form are correlated with S and show that the majority of changes in pollination mode involved a change from biotic to abiotic pollination with an associated fall in speciation rate. The majority of growth form changes involved the evolution of herbaceousness from woodiness with a correlated rise in speciation rate. We test the hypothesis of Ricklefs and Renner (1994) that “evolutionary flexibility” rather than other trait changes triggered increased speciation rates in some families, but find little support for the hypothesis.     

PHYLOGENETIC RELATIONSHIPS AMONG ISOLATES OF EUDORINA SPECIES (VOLVOCALES, CHLOROPHYTA) INFERRED FROM MOLECULAR AND BIOCHEMICAL DATA

Phylogenetic analyses of 19 strains representing five species of Eudorina, one strain of Pleodorina indica, and seven strains of Yamagishiella unicocca were carried out by sequencing the internal transcribed spacer region (ITS 1 and ITS 2) of the nuclear ribosomal DNA (rDNA) repeats. The sequence data resolved five phylogenetic groups, one consisting of Y. unicocca and the other four encompassing all the Eudorina species. Two isolates, Eudorina sp. (ASW 05157) and Pleodorina indica (ASW 05153), were of uncertain affiliation. Whereas one monophyletic group included strains of E. elegans only, the other strains of E. elegans appeared alongside E. cylindrica, E. illinoisensis, and E. unicocca var. unicocca in the other Eudorina clades. The distribution pattern of the carotenoid loroxanthin ([3R,3'R,6'R]-β,ε-carotene-3,19,3'-triol), a systematically useful biochemical marker within chlorophycean flagellates, was shown to match the evaluated molecular data. Whereas it was either totally absent or universally present in six of the deduced phylogenetic lines, it occurred randomly in the E. elegans clade containing only E. elegans isolates. The results substantiated the current hypothesis that the unique vegetative morphology of E. elegans has independently arisen at various times during evolution and that it is not a marker of a monophyletic group.     

中国蝗总科昆虫科间系统发育关系支序分析：直翅目;蝗亚目

在形态学研究的基础上,配方选择了１２个特征,利用Ｈｅｎｎｉｇ８６程序包,以支序分析探讨了中国蝗总科昆虫科间的系统发育关系。结果表明蝗虫类昆虫稳定的分为两大类,即癞蝗类和蝗类,本文建议蝗虫类昆虫分为分为两个总科较宜。长腹蝗亚科似归入斑腿蝗亚科更为合理;而皱腹亚科则应提升为独立科。     

A STUDY ON PHYLOGENETIC RELATIONSHIPS AMONG THE SUB-GENERA OF BUMBLE BEES (HYMENOPTERA:APIDAE)

Abstract Phylogenetic relationships among the 28 subgenera of bumble bees were studied by using morphological and behavioral characters. The results suggest that the traditional Bombus Latreille and Psithyrus Lepeletier are monophyletic groups. However, the position of Mendacibombus in cladograms is different from that of the traditional view. In addition, according to their geographical distribution and relative relationships Eurasian—Asian group and Eurasian—American group are recognized for Bombus Latreille.     

缘蝽科族级单元系统发育关系支序分析(半翅目:异翅亚目:缘蝽总科)

在已有比较形态学研究基础上,本文选择了15个特征方面（外生殖器为主）的74个新征,以支序分析方法探讨了缘蝽科族（或亚科）的系统发育关系。结果表明棒缘蝽亚科、希缘蝽族、沟缘蝽族较为原始,与其余类群差异较大,缘蝽科的范围值得进一步研究;除Chariesterini外的缘蝽亚科是高等的缘蝽类群;Chariesterini似应从缘蝽亚科中独立出来;狭义巨缘蝽族以及萧的鼻缘蝽族、梭缘蝽族、昧缘蝽族、曼缘蝽族成立;拟黛缘蝽属、副黛缘蝽属、华黛缘蝽属、异黛缘蝽属应从”黛缘蝽族”中分别独立出来成立新族,即拟黛缘蝽族、副黛缘蝽族、华黛缘蝽族和异黛缘蝽族,以使原有的黛缘蝽族成为自然类群。     

熊蜂族系统发育关系的研究(膜翅目:蜜蜂总科)(英文)

本文选取熊蜂雌雄外部形态特征、行为学特征,对熊蜂28个亚属之间的系统发育关系利用支序分析的方法进行了探讨。研究结果表明传统的熊蜂属与拟熊蜂属是两个单系群,Mendacibombus亚属在支序图中的位置与传统看法有所不同。另外,根据能蜂各亚属间的亲缘关系及它们的地理分布建议熊蜂属中分两个亚属群：欧亚—亚洲群与欧亚—美洲群。     

A PRELIMINARY INVESTIGATION OF THE PHYLOGENETIC RELATIONSHIPS AMONG THE FRESHWATER,APICAL PORE FIELD-BEARING CYMBELLOID AND GOMPHONEMOID DIATOMS (BACILLARIOPHYCEAE)1

Relationships among the apical pore field-bearing diatom genera Brebissonia, Cymbella, Gomphonema, Gomphoneis, Didymosphenia, Gomphocymbella, and Reimeria and their close ally Encyonema were evaluated using cladistics. These biraphid genera are a polythetic group defined by chloroplast number and type, mode of sexual reproduction, valve orientation and presence of apical pore fields. Character states of valve as well as non-siliceous features were described and then polarized using the outgroup method, with the naviculoid genera Anomoeoneis and Placoneis serving as outgroups. The cladistic analysis suggests two groups of terminal taxa, corresponding to to a predominantly cymbelloid lineage and a gomphonemoid lineage. Didymosphenia, previously thought to be closely allied to gomphonemoid diatoms, is shown to be more closely related to Cymbella. Gomphonema appears to be without distinguishing characteristics. Relationships of the other genera are described. The systematic position of small, doubly-punctate Gomphonema species is discussed.     

ORIGINS,EVOLUTION, AND DIVERSIFICATION OF CLEPTOPARASITIC LINEAGES IN LONG‐TONGUED BEES

The evolution of parasitic behavior may catalyze the exploitation of new ecological niches yet also binds the fate of a parasite to that of its host. It is thus not clear whether evolutionary transitions from free‐living organism to parasite lead to increased or decreased rates of diversification. We explore the evolution of brood parasitism in long‐tongued bees and find decreased rates of diversification in eight of 10 brood parasitic clades. We propose a pathway for the evolution of brood parasitic strategy and find that a strategy in which a closed host nest cell is parasitized and the host offspring is killed by the adult parasite represents an obligate first step in the appearance of a brood parasitic lineage; this ultimately evolves into a strategy in which an open host cell is parasitized and the host offspring is killed by a specialized larval instar. The transition to parasitizing open nest cells expanded the range of potential hosts for brood parasitic bees and played a fundamental role in the patterns of diversification seen in brood parasitic clades. We address the prevalence of brood parasitic lineages in certain families of bees and examine the evolution of brood parasitism in other groups of organisms.     

THE EVOLUTION OF DIAPAUSE IN THE KILLIFISH FAMILY RIVULIDAE (ATHERINOMORPHA,CYPRINODONTIFORMES): A MOLECULAR PHYLOGENETIC AND BIOGEOGRAPHIC PERSPECTIVE

Phylogenetic relationships within the family Rivulidae (order Cyprinodontiformes) are investigated using 1972 aligned base pairs of mitochondrial DNA (mtDNA) for samples representing 66 species. Genes analyzed include those encoding the 12S ribosomal RNA; transfer RNAs for valine, glutamine, methionine, tryptophan, alanine, asparagine, cysteine, and tyrosine; complete NADH dehydrogenase subunit II; and part of cytochrome oxidase I. Parsimony analysis of the aligned mtDNA sequences results in a single most parsimonious tree. The phylogeny reveals two independent origins of developmental diapause within the family Rivulidae. It is unlikely that diapause evolved de novo in each group, suggesting that the presence or absence of diapause is the result of developmental switches between alternative stabilized pathways. Phylogeny of the family Rivulidae shows high concordance with predictions derived from the geological history of South America and Central America. Basal lineages in the rivulid phylogeny are distributed primarily on geologically old areas, whereas more nested lineages occur in geologically younger areas. However, there is little concordance between the molecular phylogeny and currently available morphological hypotheses and existing taxonomies. Based on the mtDNA phylogeny, the genera Pterolebias, Rivulus, Pituna, and Plesiolebias are considered nonmonophyletic and warrant taxonomic reassessment.     

竹类害虫的六种寄生蜂及二新种记述（膜翅目：跳小峰科）

本文报导与竹类害虫有关的６种跳小蜂。长缘刷盾跳小蜂Ｃｈｅｉｌｏｎｅｕｒｕｓｃｌａｖｉｇｅｒ及优赛跳小蜂Ｅｕｓｅｍｉｏｎｃｏｒｎｉｇｅｒｕｍ,寄生竹巢粉蚧Ｎｅｓｔｉｃｏｃｃｕｓｓｉｎｅｎｓｉｓ的初寄生者花翅跳小蜂Ｍｉｃｒｏｔｅｒｙｓｓｐｐ．;瓢虫隐尾跳小蜂Ｈｏｍａｌｏｔｙｌｕｓｆｌａｍｉｎｉｕｓ,寄生黑缘光瓢虫Ｅｘｏｃｈｏｍｕｓ（Ｘａｎｔｈｏｃｏｒｎｓ）ｎｉｇｒｏｍａｒｇｉｎａｔｕｓ;白胫短缘跳小蜂Ｚａｏｍｍａｅｒｉｏｃｏｃｉ（Ｔａｃｈｉｋａｗａ）中国新记录,寄生竹巢粉蚧的初寄生者花翅跳小蜂;四斑长索跳小蜂ＡｎａｇｙｒｕｓｑｕａｄｒｉｍａｃｕｌａｔｕｓＸｕｅｔＨｅ,ｓｐ．ｎｏｖ,寄生竹绒粉蚧Ｅｒｉｏｃｏｃｃｕｓｔｒａｎｓｖｅｒｓｕｓ;长脉卵跳小蜂ＯｏｅｎｃｙｒｔｕｓｌｏｎｇｉｖｅｎｏｓｕｓＸｕｅｔＨｅ,ｓｐ．ｎｏｖ,寄生竹卵圆蝽Ｈｉｐｐｏｔｉｓｃｕｓｄｏｒｓａｌｉｓ。标本及模式标本保存在浙江农业大学寄生蜂标本室。     

GENETIC DIVERSITY AND INTROGRESSION IN TWO CULTIVATED SPECIES (PORPHYRA YEZOENSIS AND PORPHYRA TENERA) AND CLOSELY RELATED WILD SPECIES OF PORPHYRA (BANGIALES,RHODOPHYTA)1

We investigated the genetic variations of the samples that were tentatively identified as two cultivated Porphyra species (Porphyra yezoensis Ueda and Porphyra tenera Kjellm.) from various natural populations in Japan using molecular analyses of plastid and nuclear DNA. From PCR‐RFLP analyses using nuclear internal transcribed spacer (ITS) rDNA and plastid RUBISCO spacer regions and phylogenetic analyses using plastid rbcL and nuclear ITS‐1 rDNA sequences, our samples from natural populations of P. yezoensis and P. tenera showed remarkably higher genetic variations than found in strains that are currently used for cultivation. In addition, it is inferred that our samples contain four wild Porphyra species, and that three of the four species, containing Porphyra kinositae, are closely related to cultivated Porphyra species. Furthermore, our PCR‐RFLP and molecular phylogenetic analyses using both the nuclear and plastid DNA demonstrated the occurrence of plastid introgression from P. yezoensis to P. tenera and suggested the possibility of plastid introgression from cultivated P. yezoensis to wild P. yezoensis. These results imply the importance of collecting and establishing more strains of cultivated Porphyra species and related wild species from natural populations as genetic resources for further improvement of cultivated Porphyra strains.     

天牛科昆虫高级分类阶元实体的进化研究

本文从生物地理学、形态学、生态学、行为学、生理学、天牛—寄主植物的相互关系、地质学、气候变迁等诸方面,探讨了天牛科高级分类阶元实体的进化证据,提出了各阶元实体进化关系的新假说。研究结果表明,沟胫天牛亚科、锯天牛亚科和天牛亚科可能于侏罗纪在冈瓦纳古陆上分化出来;花天牛亚科和幽天牛亚科于白垩纪在劳亚古陆上分化出来。各亚科的进化顺序有两种可能:(1)沟胫天牛亚科、锯天牛亚科、天牛亚科、花天牛亚科、幽天牛亚科;(2)沟胫天牛亚科、天牛亚科、锯天牛亚科;花天牛亚科、幽天牛亚科。文章最后还讨论了各亚科的系统发育关系。     

CRYPTIC DIVERSITY AND PHYLOGENETIC RELATIONSHIPS WITHIN THE MASTOCARPUS PAPILLATUS SPECIES COMPLEX (RHODOPHYTA,PHYLLOPHORACEAE)1

Mastocarpus papillatus (C. Agardh) Kütz. is a common intertidal red alga occurring along the west coast of North America from Baja California to Alaska. Sequencing of both the chloroplast‐encoded rbcL gene and the nuclear ribosomal internal transcribed spacer (ITS) regions of ～200 specimens from California to Alaska revealed that M. papillatus is actually a complex of at least five species. All five species have high bootstrap support in phylogenetic analyses of both genetic regions, and in the case of the ITS marker, the species also have distinctive patterns of indels. Three of the species are localized in the mid‐ to upper intertidal, whereas two of the species occur in the low intertidal. The species also have different geographic ranges that overlap in the Vancouver Island area of British Columbia. No distinctive, reliable morphological differences were observed among the species. Although a variety of names are available for species in the complex, it is not yet clear which name goes with which species. As part of the survey, I also sequenced other species of Mastocarpus in the northeast Pacific region, and I provide new distribution records for M. jardinii ( J. Agardh) J. A. West and for a nonpapillate and probably undescribed species of Mastocarpus.     

MAJOR DEVELOPMENTAL REGULATORS AND THEIR EXPRESSION IN TWO CLOSELY RELATED SPECIES OF PORPHYRA (RHODOPHYTA)1

Little is known about the genetic and biochemical mechanisms that underlie red algal development, for example, why the group failed to evolve complex parenchyma and tissue differentiation. Here we examined expressed sequence tag (EST) data from two closely related species, Porphyra umbilicalis (L.) J. Agardh and P. purpurea (Roth) C. Agardh, for conserved developmental regulators known from model eukaryotes, and their expression levels in several developmental stages. Genes for most major developmental families were present, including MADS‐box and homeodomain (HD) proteins, SNF2 chromatin‐remodelers, and proteins involved in sRNA biogenesis. Some of these genes displayed altered expression correlating with different life history stages or cell types. Notably, two ESTs encoding HD proteins showed eightfold higher expression in the P. purpurea sporophyte (conchocelis) than in the gametophyte (blade), whereas two MADS domain‐containing paralogs showed significantly different patterns of expression in the conchocelis and blade respectively. These developmental gene families do not appear to have undergone the kinds of dramatic expansions in copy number found in multicellular land plants and animals, which are important for regulating developmental processes in those groups. Analyses of small RNAs did not validate the presence of miRNAs, but homologs of Argonaute were present. In general, it appears that red algae began with a similar molecular toolkit for directing development as did other multicellular eukaryotes, but probably evolved altered roles for many key proteins, as well as novel mechanisms yet to be discovered.     

PIGMENT SIGNATURES AND PHYLOGENETIC RELATIONSHIPS OF THE PAVLOVOPHYCEAE (HAPTOPHYTA)1

Variations in the HPLC‐derived pigment composition of cultured Pavlovophyceae (Cavalier‐Smith) Green et Medlin were compared with phylogenetic relationships inferred from 18S rDNA sequencing, morphological characteristics, and current taxonomy. The four genera described for this haptophyte class (Diacronema Prauser emend. Green et Hibberd, Exanthemachrysis Lepailleur, Pavlova Butcher, and Rebecca Green) were represented by nine different species (one of which with data from GeneBank only). Chlorophylls a, c₁, c₂ and MgDVP (Mg‐[3,8‐divinyl]‐phytoporphyrin‐13²‐methylcarboxylate) and the carotenoids fucoxanthin, diadinoxanthin, diatoxanthin, and β,β‐carotene were detected in all cultures. Species only differed in the content of an unknown (diadinoxanthin‐like) xanthophyll and two polar chl c forms, identified as a monovinyl (chl c₁‐like) and a divinyl (chl c₂‐like) compound. This is the first observation of the monovinyl form in haptophytes. Based on distribution of these two chl c forms, species were separated into Pavlovophyceae pigment types A, B, and C. These pigment types crossed taxonomic boundaries at the generic level but were in complete accordance with species groupings based on molecular phylogenetic relationships and certain ultrastructural characteristics (position and nature of pyrenoid, stigma, and flagella). These results suggest that characterization of the pigment signature of unidentified culture strains of Pavlovophyceae can be used to predict their phylogenetic affinities and vice versa. Additional studies have been initiated to evaluate this possibility for the haptophyte class Prymnesiophyceae.     

基于线粒体Cyt b基因和CO I基因序列研究豹蛱蝶亚科(鳞翅目,蛱蝶科)10属间的系统发生关系

测定了国产豹蛱蝶亚科10属共10个代表种的Cyt 6基因和CO Ⅰ基因的部分序列.结合从GenBank中获得的3个种类CO Ⅰ基因的同源序列,以锯眼蝶亚科2个物种为外群,通过遗传分析软件对CO Ⅰ、Cyt b独立基因序列和联合基因序列以及编码的氨基酸序列进行了比较分析,同时用邻接法(NJ)、最大简约法(MP)、最大似然法(ML)和贝叶斯法(BI)重建分子系统树,分析了该亚科10个属之间的系统发生关系.结果显示:1)联合基因序列的A+T平均含量为71.90h,具A、T偏倚性,其编码的357个氨基酸中没有半胱氨酸,变异率为11.5%;2)豹蛱蝶亚族为单系群;3)青豹蛱蝶属和豹蛱蝶属间、黄襟蛱蝶属和珐蛱蝶属间具有较近的亲缘关系;4)支持将文蛱蝶属、襟蛱蝶属和珐蛱蝶属从豹蛱蝶亚科中分离出来.     

CONVERGENT EVOLUTION OF COURTSHIP SONGS AMONG CRYPTIC SPECIES OF THE CARNEA GROUP OF GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE: CHRYSOPERLA)

Although traits of related species are likely to be similar due to common ancestry, mating signals are an exception. In singing insects, for example, song similarity has been documented only for allopatric or allochronic species pairs, and even then, not often. Where song similarity does occur, it has been logically attributed to the inheritance of ancestral traits rather than convergence. It is quite common for related, sympatric insect species to differ dramatically in calling song, which is predicted by evolutionary theory to maximize intraspecific mating success. Given that there are a limited number of ways to make sounds on anatomically similar organs and given that there would be no selective pressure for songs to differ in widely separated geographic areas, convergence in songs among related species living on different continents might be expected. Here we present the first well-documented case of such convergence, in a group of sibling, cryptic species characterized by substrate-borne vibrational mating songs. In this example from green lacewings of the carnea group of the genus Chrysoperla, a variety of statistical tests shows that one species in North America and another in Asia possess songs that are strikingly similar to each other. DNA data demonstrate that the species involved belong to divergent speciose lineages, and behavioral data demonstrate that the convergent songs are readily accepted by members of both species.