Evolutionary relationships of sibling tapeworm species (Cestoda) parasitizing teleost fishes

DNA/DNA hybridization and sequencing of rDNA (partial 18S rDNA and ITS1) were used to investigate phylogenetic relationships among seven host-specific Bothriocephalus parasites (Cestoda, Pseudophyllidae). The small nucleotide divergence between six of the seven bothriocephalids suggests that isolation and differentiation of Bothriocephalus lineages in the different host species probably occurred recently and over a short time span. Comparison of the molecular phylogeny of the parasite species to the phylogeny of their hosts (teleostean fishes) revealed little congruence between the branching patterns of hosts and parasites, suggesting that bothriocephalids have not cospeciated with their hosts.      

Native saltbush (Rhagodia spp.; Chenopodiaceae) as a potential reservoir for agromyzid leafminer parasitoids on horticultural farms

Australia has to date been spared the introduction of highly polyphagous invasive pest agromyzid leafminers; however, their arrival and spread should be considered imminent. To develop a pre-emptive control strategy to deal with exotic leafminer outbreaks the first step is to identify Australian leafmining flies, their plant hosts and their parasitoids to gain an understanding of their population dynamics. Native vegetation may be providing resources for beneficial parasitic wasps plus access to alternative hosts and refuge from disturbance. Here, two Australian endemic saltbushes ( Rhagodia candolleana and R. parabolica , Caryophyllales: Chenopodiaceae) have been investigated for their potential to act as reservoirs for endemic agromyzid hosts and their key parasitoids. Mined leaves of the two Rhagodia species were sampled on two commercial horticultural properties in the Virginia horticulture area on the Northern Adelaide Plains between September 2007 and April 2008. Leaf mines on both Rhagodia species were caused by an endemic leafminer species, putatively Phytoliriomyza praecellens Spencer (Diptera: Agromyzidae). Ten species of parasitoids (all Hymenoptera) emerged from R. candolleana mines and seven different species from R. parabolica mines, mainly from the family Eulophidae and with some Pteromalidae and Braconidae. Trigonogastrella Girault sp. (Pteromalidae), Zagrammosoma latilineatum Ubaidillah and Hemiptarsenus varicornis Girault (both Eulophidae) were the most abundant species on R. candolleana , whereas two Opius Wesmael spp. (Braconidae) were the most abundant species on R. parabolica . Findings from this survey suggest an opportunity to plant purpose-designed refuges that could play a role in conservation biological control as part of an Integrated Pest Management strategy developed prior to incursion of pest leafminers such as Liriomyza species.     

Molecular phylogeny of gill monogeneans (Platyhelminthes, Monogenea, Dactylogyridae) and colonization of Indo-West Pacific butterflyfish hosts (Perciformes, Chaetodontidae)

We investigated the phylogenetic relationships among monogenean parasites of the Chaetodontidae (butterflyfishes) from the Indo-West Pacific Ocean. Molecular phylogenies of selected taxa within the Dactylogyridae, including the ancyrocephaline parasites of butterflyfishes, based on two nuclear and one mitochondrial gene fragments (complete 18S rDNA, partial 28S rDNA (D1-D3), and partial 16S rDNA) were reconstructed using parsimony, maximum likelihood and Bayesian inference methods. Our results show the non-monophyletic nature of the monogenean fauna of butterflyfishes. The group is divided into two independent lineages. The first clade contains species of the genera Aliatrema and Euryhaliotrematoides , which parasitize Chaetodontidae exclusively. The second contains species of Haliotrema , a generalist group of parasites. The positions of several other species of the Ancyrocephalinae, including freshwater species, at the base of the two clades, provide strong evidence that the monogenean fauna did not result from a single colonization event, but rather that they have colonized their butterflyfish hosts independently at least twice.     

Evidence for cospeciation events in the host–symbiont system involving crinoids (Echinodermata) and their obligate associates,the myzostomids (Myzostomida,Annelida)

Although molecular-based phylogenetic studies of hosts and their associates are increasingly common in the literature, no study to date has examined the hypothesis of coevolutionary process between hosts and commensals in the marine environment. The present work investigates the phylogenetic relationships among 16 species of obligate symbiont marine worms (Myzostomida) and their echinoderm hosts (Crinoidea) in order to estimate the phylogenetic congruence existing between the two lineages. The combination of a high species diversity in myzostomids, their host specificity, their wide variety of lifestyles and body shapes, and millions years of association, raises many questions about the underlying mechanisms triggering their diversification. The phylogenetic relationships, inferred using a three-genes dataset (18S rDNA, 16S rDNA, and COI) and two-genes dataset (18S rDNA, and COI) for the myzostomids and crinoids, respectively, were congruent with the literature. The overall congruence between the two phylogenies was statistically significant according to topology-based, distance-based, and data-based approaches: a significant pattern of cophylogeny was found, though not perfect probably resulting from occasional host switches, duplications or extinction events. A minimum of 8 cospeciation events was estimated, which is significantly higher than it would have been expected due to chance alone.     

Host discrimination by Elasmus schmitti (Hymenoptera, Eulophidae) and Latibulus argiolus (Hymenoptera, Ichneumonidae), parasitoids of colonies of Polistes wasps (Hymenoptera, Vespidae)

The ability to discriminate parasitized and non-parasitized hosts allows the parasitoids to avoid intraspecific and often interspecific competition. In 2005–2009, the incidence of mixed infestation of Polistes nimpha (Christ) colonies by parasitoids Elasmus schmitti Ruschka (Hymenoptera, Eulophidae) and Latibulus argiolus (Rossi) (Hymenoptera, Ichneumonidae) was analyzed. In settlements of different types, the frequencies of mixed infestation deviated from the values expected in case of random distribution. The presence or absence of discrimination ability cannot be confirmed unequivocally based on our material. The data obtained are discussed in relation to the biological features of parasitoids.     

Taxonomic Composition of Hymenopteran Parasitoid Assemblages from Agromyzid Leaf-miners Sampled in Central Argentina

This paper deals with the taxonomic composition of parasitoid assemblages reared from agromyzid leaf-miners of 28 species collected in Cordoba, Central Argentina. The relative representation of each parasitic family was estimated, taking into consideration their species richness and abundance. Sample size affected the total species richness in each complex, and also that of Eulophidae and Pteromalidae, indicating that not only size but also taxonomic structure of parasitic assemblages are influenced by sampling effort. 3 superfamilies and 5 families of Hymenoptera Parasitica were represented, with Eulophidae (Chalcidoidea) specifically and numerically dominant both in the total community of parasitoids and in the parasitic assemblages of individual hosts. Chrysonotomyia, Chrysocharis (Eulophidae), Halticoptera (Pteromalidae) and Phaedrotoma (Braconidae) were the most relevant genera in terms of number of species and number of specimens reared. The parasitoids obtained from samples taken in agroecosystems differed from the global spectrum of taxa mainly by showing a numerical dominance of Braconidae, despite a noticeable specific impoverishment of this family.     

A survey of hymenopteran parasitoids of forest macrolepidoptera in the central Appalachians

In 1995 and 1996, we conducted a study of the hymenopteran parasitoids of macrolepidopteran larvae in the George Washington National Forest (GWNF), Augusta County, Virginia, and the Monongahela National Forest (MNF), Pocahontas County, West Virginia. Macrolepidopteran larvae were collected from canopy foliage and from under canvas bands placed around tree boles. A total of 115 macrolepidopteran species and 5,235 individual larvae were reared. Forty-two percent (2,221) of the larvae were gypsy moth, Lymantria dispar (L.) (Lymantriidae). A total of 43 primary and secondary (hyperparasitoid) hymenopteran parasitoid species were reared from 46 macrolepidopteran species. Hymenopteran families represented included Ichneumonidae (23 species), Braconidae (19), Eulophidae (6), Perilampidae (1), and Trigonalidae (1). We reared 41 and 28 parasitoid species from the GWNF and the MNF, respectively, with 19 species reared from both forests. Many parasitoid species were collected infrequently, suggesting that they are relatively rare on the sampled hosts. The introduced species Cotesia melanoscela (Ratzeburg) (Braconidae), and Euplectrus bicolor (Swederus) (Eulophidae) were among the most commonly reared parasitoids, the latter reared from native hosts. The four most commonly reared native parasitoids were Meteorus hyphantriae, Riley (Braconidae), Microplitis near hyphantriae (Ashmead) (Braconidae), Aleiodes preclarus Marsh & Shaw, and Euplectrus maculiventris (Westwood) (Eulophidae). A total of 53 new hymenopteran parasitoid-macrolepidopteran host records were documented. Results from this study will be used to evaluate long-term treatment effects of regional applications of Bacillus thuringiensis kurstaki, and the gypsy moth fungus Entomophaga maimaiga Humber, Shimazu & Soper on hymenopteran parasitoids of macrolepidopteran larvae.     

Effects of sequence alignment and structural domains of ribosomal DNA on phylogeny reconstruction for the protozoan family sarcocystidae

Finding correct species relationships using phylogeny reconstruction based on molecular data is dependent on several empirical and technical factors. These include the choice of DNA sequence from which phylogeny is to be inferred, the establishment of character homology within a sequence alignment, and the phylogeny algorithm used. Nevertheless, sequencing and phylogeny tools provide a way of testing certain hypotheses regarding the relationship among the organisms for which phenotypic characters demonstrate conflicting evolutionary information. The protozoan family Sarcocystidae is one such group for which molecular data have been applied phylogenetically to resolve questionable relationships. However, analyses carried out to date, particularly based on small-subunit ribosomal DNA, have not resolved all of the relationships within this family. Analysis of more than one gene is necessary in order to obtain a robust species signal, and some DNA sequences may not be appropriate in terms of their phylogenetic information content. With this in mind, we tested the informativeness of our chosen molecule, the large-subunit ribosomal DNA (lsu rDNA), by using subdivisions of the sequence in phylogenetic analysis through PAUP, fastDNAml, and neighbor joining. The segments of sequence applied correspond to areas of higher nucleotide variation in a secondary-structure alignment involving 21 taxa. We found that subdivision of the entire lsu rDNA is inappropriate for phylogenetic analysis of the Sarcocystidae. There are limited informative nucleotide sites in the lsu rDNA for certain clades, such as the one encompassing the subfamily Toxoplasmatinae. Consequently, the removal of any segment of the alignment compromises the final tree topology. We also tested the effect of using two different alignment procedures (CLUSTAL W and the structure alignment using DCSE) and three different tree-building methods on the final tree topology. This work shows that congruence between different methods in the formation of clades may be a feature of robust topology; however, a sequence alignment based on primary structure may not be comparing homologous nucleotides even though the expected topology is obtained. Our results support previous findings showing the paraphyly of the current genera Sarcocystis and Hammondia and again bring to question the relationships of Sarcocystis muris, Isospora felis, and Neospora caninum. In addition, results based on phylogenetic analysis of the structure alignment suggest that Sarcocystis zamani and Sarcocystis singaporensis, which have reptilian definitive hosts, are monophyletic with Sarcocystis species using mammalian definitive hosts if the genus Frenkelia is synonymized with Sarcocystis.     

Phylogeny of the glomeromycota (arbuscular mycorrhizal fungi): recent developments and new gene markers

The fungal symbionts of arbuscular mycorrhiza form a monophyletic group in the true Fungi, the phylum Glomeromycota. Fewer than 200 described species currently are included in this group. The only member of this clade known to form a different type of symbiosis is Geosiphon pyriformis, which associates with cyanobacteria. Because none of these fungi has been cultivated without their plant hosts or cyanobacterial partners, progress in obtaining multigene phylogenies has been slow and the nuclear-encoded ribosomal RNA genes have remained the only widely accessible molecular markers. rDNA phylogenies have revealed considerable polyphyly of some glomeromycotan genera that has been used to reassess taxonomic concepts. Environmental studies using phylogenetic methods for molecular identification have recovered an amazing diversity of unknown phylotypes, suggesting considerable cryptic species diversity. Protein gene sequences that have become available recently have challenged the rDNA-supported sister group relationship of the Glomeromycota with Asco/Basidiomycota. However the number of taxa analyzed with these new markers is still too small to provide a comprehensive picture of intraphylum relationships. We use nuclear-encoded rDNA and rpb1 protein gene sequences to reassess the phylogeny of the Glomeromycota and discuss possible implications.     

Mitochondrial phylogeography of a leafminer parasitoid, Diglyphus isaea (Hymenoptera: Eulophidae) in China

Diglyphus isaea is one of the primary parasitoids of agromyzid leafminers. Mitochondrial cytochrome oxidase subunit I sequences from 10 Chinese localities were used to investigate geographic variation within this widespread and morphologically uniform parasitoid. Phylogeographic analyses and sequence variation indicated that this species was actually a species-complex. We used an amber fossil record of the genus Entedon (Eulophidae: Entedoninae) as a reference to estimate the divergence time of major clades within D. isaea. With reference to the fossil record of its hosts, our results indicate that associations between parasitoids and leaf-miners could have been possibly established 38.2 million years ago.     

Abundance,not diversity,of host beetle communities determines abundance and diversity of parasitoids in deadwood

Most parasites and parasitoids are adapted to overcome defense mechanisms of their specific hosts and hence colonize a narrow range of host species. Accordingly, an increase in host functional or phylogenetic dissimilarity is expected to increase the species diversity of parasitoids. However, the local diversity of parasitoids may be driven by the accessibility and detectability of hosts, both increasing with increasing host abundance. Yet, the relative importance of these two mechanisms remains unclear. We parallelly reared communities of saproxylic beetle as potential hosts and associated parasitoid Hymenoptera from experimentally felled trees. The dissimilarity of beetle communities was inferred from distances in seven functional traits and from their evolutionary ancestry. We tested the effect of host abundance, species richness, functional, and phylogenetic dissimilarities on the abundance, species richness, and Shannon diversity of parasitoids. Our results showed an increase of abundance, species richness, and Shannon diversity of parasitoids with increasing beetle abundance. Additionally, abundance of parasitoids increased with increasing species richness of beetles. However, functional and phylogenetic dissimilarity showed no effect on the diversity of parasitoids. Our results suggest that the local diversity of parasitoids, of ephemeral and hidden resources like saproxylic beetles, is highest when resources are abundant and thereby detectable and accessible. Hence, in some cases, resources do not need to be diverse to promote parasitoid diversity.     

Robustness of ancestral state estimates: evolution of life history strategy in ichneumonoid parasitoids

We test hypotheses for the evolution of a life history trait among a group of parasitoid wasps (Hymenoptera: Ichneumonoidea), namely, the transition among koinobiont parasitoids (parasitoids whose hosts continue development after oviposition) between attacking exposed hosts and attacking hosts that are concealed within plant tissue. Using a range of phylogeny estimates based on 28S rDNA sequences, we use maximum parsimony (MP) and maximum likelihood (ML) methods to estimate the ancestral life history traits for the main clades in which both traits occur (using the programs MacClade and Discrete, respectively). We also assess the robustness of these estimates; for MP, we use step matrices in PAUP* to find the minimum weight necessary to reverse estimates or make them ambiguous, and for ML, we measure the differences in likelihood after fixing the ancestral nodes at the alternative states. We also measure the robustness of the MP ancestral state estimate against uncertainties in the phylogeny estimate, manipulating the most-parsimonious tree in MacClade to find the shortest suboptimal tree in which the ancestral state estimate is reversed or made ambiguous. Using these methods, we find strong evidence supporting two transitions among koinobiont Ichneumonoidea: (1) to attacking exposed hosts in a clade consisting of the Helconinae and related subfamilies, and (2) the reverse transition in a clade consisting of the Euphorinae and related subfamilies. In exploring different methods of analyzing variable-length DNA sequences, we found that direct optimization with POY gave some clearly erroneous results that had a profound effect on the overall phylogeny estimate. We also discuss relationships within the superfamily and expand the Mesostoinae to include all the gall-associated braconids that form the sister group of the Aphidiinae.     

基于28S rDNA D2基因序列的几种寄生夜蛾科的多胚跳小蜂的分子系统发育（英文）

本文基于28S rDNA D2部分序列, 用MP和ML方法对几种寄生夜蛾科的多胚跳小蜂Copidosoma spp.进行了系统发育研究,并对其形态和生物学关系进行了讨论。结果表明：寄生金翅夜蛾亚科(Plusiinae)的C. floridanum 和寄生实夜蛾亚科(Heliothinae)的C. primulum关系较近,而以夜蛾亚科(Noctuinae)为寄主的C. truncatellum 和 C. agrotis关系更近。结果提示核基因28S D2区可能对研究多胚跳小蜂种间的系统发育关系很有帮助。     

The Evolution of Body Size,Antennal Size and Host Use in Parasitoid Wasps (Hymenoptera: Chalcidoidea): A Phylogenetic Comparative Analysis

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.     

基于18S基因序列的姬小蜂分子系统发育

本文基于18S rDNA部分序列,用MP和Baysian方法研究了姬小蜂科的系统发育,对姬小蜂科的单系性及其与其它小蜂科间的关系进行了讨论。姬小蜂亚科、灿姬小蜂亚科和啮姬小蜂亚科形成三个独立的支系,研究结果支持它们各自的单系性,但本结果没有明确姬小蜂科的单系性。研究结果同时还支持瑟姬小蜂族、扁股姬小蜂族和狭面姬小蜂族三个族的地位,但不支持姬小蜂族的地位。姬小蜂科的单系性及其与其它小蜂间的关系还需更多的形态学数据和更多的基因序列来进一步研究[动物学报52 (2) : 288 -301 , 2006]。     

The roles of ecological fitting, phylogeny and physiological equivalence in understanding realized and fundamental host ranges in endoparasitoid wasps

Co-evolutionary theory underpins our understanding of interactions in nature involving plant-herbivore and host-parasite interactions. However, many studies that are published in the empirical literature that have explored life history and development strategies between endoparasitoid wasps and their hosts are based on species that have no evolutionary history with one another. Here, we investigated novel associations involving two closely related solitary endoparasitoids that originate from Europe and North America and several of their natural and factitious hosts from both continents. The natural hosts of both species are also closely related, all being members of the same family. We compared development and survival of both parasitoids on the four host species and predicted that parasitoid performance is better on their own natural hosts. In contrast with this expectation, survival, adult size and development time of both parasitoids were similar on all (with one exception) hosts, irrespective as to their geographic origin. Our results show that phylogenetic affinity among the natural and factitious hosts plays an important role in their nutritional suitability for related parasitoids. Evolved traits in parasitoids, such as immune suppression and development, thus enable them to successfully develop in novel host species with which they have no evolutionary history. Our results show that host suitability for specialized organisms like endoparasitoids is closely linked with phylogenetic history and macro-evolution as well as local adaptation and micro-evolution. We argue that the importance of novel interactions and 'ecological fitting' based on phylogeny is a greatly underappreciated concept in many resource-consumer studies.     

米蛾体内Wolbachia的wsp基因序列测定与系统发育分析

Wolbachia是广泛分布于节肢动物体内的一类共生菌, 它们参与多种调控寄主的生殖活动机制。通过对wsp基因的特异性扩增和测序,发现了Wolbachia在米蛾Corcyra cephalonica (Stainton)体内的感染。利用所测序列和其他已发表的序列建立系统树,结果表明米蛾体内Wolbachia属于B大组的Pip类群,与其寄生物茧蜂及赤眼蜂中的Wolbachia各株系遗传距离相差较远。据此推测米蛾体内感染的Wolbachia不是由寄生物（茧蜂、赤眼蜂）水平传播所致。     

Piloting the use of DNA barcoding in support of natural enemy surveys: New parasitoid records for banana skippers (Erionota spp., Lepidoptera,Hesperiidae) in Malaysia

We report a preliminary study carried out in Peninsular Malaysia, with the objective of rearing parasitoids of the two species of banana skippers (Erionota thrax and E. torus: Lepidoptera, Hesperiidae) and using DNA barcoding of Erionota spp. early stages (which are indistinguishable) and host remains to associate parasitoids with their host Erionota species. We were able to use barcodes to reliably identify the dead host remains, unreliably identify a limited subset of associated material (exuviae, frass, pupal exoskeleton, silk, wax) and to make provisional generic identifications of the parasitoids. We found parasitism by Argyrophylax sp. possibly phoeda (Tachinidae) on both species of Erionotus, by ?Casinaria sp. (Ichneumonidae) on E. thrax, and by Elasmus sp. (Eulophidae) on E. torus. The first is a new host-parasitoid association, and the second is a new association for Peninsular Malaysia. We conclude that the use of DNA barcoding, particularly for recently dead host remains, is a valuable tool to assess host-parasitoid associations where the hosts cannot be reliably identified from morphological characters.     

Role of emerald ash borer (Coleoptera: Buprestidae) larval vibrations in host-quality assessment by Tetrastichus planipennisi (Hymenoptera: Eulophidae)

The biological control agent Tetrastichus planipennisi Yang (Hymenoptera: Eulophidae) is a gregarious larval endoparasitoid of the emerald ash borer, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), an invasive cambium-feeding species responsible for recent, widespread mortality of ash (Fraxinus spp.) in North America. T. planipennisi is known to prefer late-instar emerald ash borer, but the cues used to assess host size by this species and most other parasitoids of concealed hosts remain unknown. We sought to test whether vibrations produced by feeding emerald ash borer vary with larval size and whether there are any correlations between these cues and T. planipennisi progeny number (i.e., brood size) and sex ratio. The amplitudes and rates of 3-30-ms vibrational impulses produced by emerald ash borer larvae of various sizes were measured in the laboratory before presenting the larvae to T. planipennisi. Impulse-rate did not vary with emerald ash borer size, but vibration amplitude was significantly higher for large larvae than for small larvae. T. planipennisi produced a significantly higher proportion of female offspring from large hosts than small hosts and was shown in previous work to produce more offspring overall from large hosts. There were no significant correlations, however, between the T. planipennisi progeny data and the emerald ash borer sound data. Because vibration amplitude varied significantly with host size, however, we are unable to entirely reject the hypothesis that T. planipennisi and possibly other parasitoids of concealed hosts use vibrational cues to assess host quality, particularly given the low explanatory potential of other external cues. Internal chemical cues also may be important.     

伊朗粉虱寄生蜂种类记述(膜翅目:广腹细蜂科、姬小蜂科、蚜小蜂科)

本文报道了伊朗寄生粉虱的25种寄生蜂,分别隶属于膜翅目的广腹细蜂科、姬小蜂科和蚜小蜂科,其中15种寄生蜂种类为伊朗的分布新纪录(标有*号者)。