Alejandro Zaldívar-Riverón

Molecular phylogeny and historical biogeography of the cosmopolitan parasitic wasp subfamily Doryctinae (Hymenoptera : Braconidae)

The phylogenetic relationships among representatives of 64 genera of the cosmopolitan parasitic wasps of the subfamily Doryctinae were investigated based on nuclear 28S ribosomal (r) DNA (~650 bp of the D2–3 region) and cytochrome c oxidase I (COI) mitochondrial (mt) DNA (603 bp) sequence data. The molecular dating of selected clades and the biogeography of the subfamily were also inferred. The partitioned Bayesian analyses did not recover a monophyletic Doryctinae, though the relationships involved were only weakly supported. Strong evidence was found for rejecting the monophylies of both Doryctes Haliday, 1836 and Spathius Nees, 1818. Our results also support the recognition of the Rhaconotini as a valid tribe. A dispersal–vicariance analysis showed a strong geographical signal for the taxa included, with molecular dating estimates for the origin of Doryctinae and its subsequent radiation both occurring during the late Paleocene–early Eocene. The divergence time estimates suggest that diversification in the subfamily could have in part occurred as a result of continental break-up events that took place in the southern hemisphere, though more recent dispersal events account for the current distribution of several widespread taxa.

DNA barcoding and the taxonomy of Microgastrinae wasps (Hymenoptera, Braconidae): Impacts after 8 years and nearly 20 000 sequences

Microgastrine wasps are among the most species‐rich and numerous parasitoids of caterpillars (Lepidoptera). They are often host‐specific and thus are extensively used in biological control efforts and figure prominently in trophic webs. However, their extraordinary diversity coupled with the occurrence of many cryptic species produces a significant taxonomic impediment. We present and release the results of 8 years (2004–2011) of DNA barcoding microgastrine wasps. Currently they are the best represented group of parasitoid Hymenoptera in the Barcode of Life Data System (BOLD), a massive barcode storage and analysis data management site for the International Barcoding of Life (iBOL) program. There are records from more than 20 000 specimens from 75 countries, including 50 genera (90% of the known total) and more than 1700 species (as indicated by Barcode Index Numbers and 2% MOTU). We briefly discuss the importance of this DNA data set and its collateral information for future research in: (1) discovery of cryptic species and description of new taxa; (2) estimating species numbers in biodiversity inventories; (3) clarification of generic boundaries; (4) biological control programmes; (5) molecular studies of host‐parasitoid biology and ecology; (6) evaluation of shifts in species distribution and phenology; and (7) fostering collaboration at national, regional and world levels. The integration of DNA barcoding with traditional morphology‐based taxonomy, host records, and other data has substantially improved the accuracy of microgastrine wasp identifications and will significantly accelerate further studies on this group of parasitoids.

Species identification in the taxonomically neglected, highly diverse, neotropical parasitoid wasp genus Notiospathius (Braconidae: Doryctinae) based on an integrative molecular and morphological approach.

Various DNA sequence-based methods for species delineation have recently been developed to assess the species-richness of highly diverse, neglected invertebrate taxa. These methods, however, need to be tested under a variety of conditions, including the use of different markers and parameters. Here, we explored the species diversity of a species-rich group of braconid parasitoid wasps, the Neotropical genus Notiospathius, including 233 specimens from 10 different countries. We examined sequences of two mitochondrial (mt) (COI, cyt b) and one nuclear (wg) gene fragments. We analysed them separately as well as concatenating the mt data with the general mixed Yule-coalescent (GMYC) model for species delineation using different tree-building methods and parameters for reconstructing ultrametric trees. We evaluated the performance of GMYC analyses by comparing their species delineations with our morphospecies identifications. Reconstructing ultrametric trees with a relaxed lognormal clock rate using the program BEAST gave the most congruent results with morphology for the two mt markers. A tree obtained with wg using the programs MrBayes+Pathd8 had the fewest cases of incongruence with morphology, though the performance of this nuclear marker was considerably lower than that of COI and cyt b. Species delimitation using the coalescent prior to obtain ultrametric trees was morphologically more congruent with COI, whereas the Yule prior was more congruent with cyt b. The analyses concatenating the mt datasets failed to recover some species supported both by morphology and the separate analyses of the mt markers. The highest morphological congruence was obtained with the GMYC analysis on an ultrametric tree reconstructed with cyt b using the relaxed lognormal clock rate and the Yule prior, thus supporting the importance of using alternative markers when the information of the barcoding locus (COI) is not concordant with morphological evidence. Seventy-one species were delimited based on the congruence found among COI, cyt b and morphology. Both mt markers also revealed the existence of seven potential cryptic species. This high species richness from a scattered geographical sampling indicates that there is a remarkable number of Notiospathius species that remains undiscovered.

Phylogeny of the Mexican coastal leopard frogs of the Rana berlandieri group based on mtDNA sequences

Phylogenetic relationships among specimens from 25 different locations for the six Mexican coastal leopard frog species of the Rana berlandieri species group were investigated using 797 bp of the mitochondrial 12S rDNA gene. Relationships among the haplotypes obtained were recovered using maximum parsimony and Bayesian analyses. Most of the clades recovered by both tree building methods are strongly supported, but conflicting clades recovered by each analysis are generally poorly supported. Both analyses reject the previously proposed subgroupings of the R. berlandieri species group. Based on the strongly supported relationships, genetic differentiation, and geographic distribution of the haplotypes examined, nine independent lineages appear to comprise the group of study. However, confirmation of the new proposed lineages will require further analyses based on other genetic markers and additional samples that cover their entire geographic distribution. Concordance was noted between Miocene-Pliocene geological and climatic events in Mexico and the relationships recovered among the lineages proposed and their geographic distribution.

A New Emerald Ash Borer (Coleoptera: Buprestidae) Parasitoid Species of Spathius Nees (Hymenoptera: Braconidae: Doryctinae) from the Russian Far East and South Korea

ABSTRACT A new emerald ash borer (Coleoptera: Buprestidae) ectoparasitoid species, Spathius galinae Belokobylskij & Strazanac (Hymenoptera: Braconidae: Doryctinae), is described from the Russian Far East and South Korea. Molecular evidence supports that Russian and Korean specimens represent a single species and are closely related to two Asian species (S. agrili and S. generosus) that belong to the S. exarator species group. The morphological variation of adult S. galinae and its immature stages, distribution, life history, and relation to similar Asian Spathius species are discussed. The distribution of the new species may indicate this is one of the more cold hardy emerald ash borer parasitoids. A brief review of hymenopteran parasitoids of emerald ash borer and new host record, the first for Atanycolus nigriventris Vojnovskaja-Krieger (Braconidae: Braconinae), is reported.

DNA barcoding a highly diverse group of parasitoid wasps (Braconidae: Doryctinae) from a Mexican nature reserve.

Background and aims. The preliminary results of a DNA barcoding study of the doryctine fauna of parasitoid wasps from the Chamela–Cuixmala Biosphere Reserve in Mexico, a region dominated by tropical dry forest, are presented. So far, three field trips have been carried out to the reserve and 468 specimens have been collected, of which 407 cox1 sequences were obtained. Materials and methods. The general mixed Yule-coalescent model was applied to a phylogram to investigate the number of evolutionary units that can be detected from the DNA sequence data examined. Results. A total of 185 barcoding species assigned to 20 identified doryctine genera were discriminated using the above model, 115 of which belong to the speciose genus Heterospilus, pointing out the extraordinary species richness of this subfamily of insects in a Mexican tropical dry forest. Conclusion. On the basis of the DNA barcodes generated, Ptesimogastroides Braet & van Achterberg is proposed to be a junior synonym of Ptesimogaster Marsh syn. nov. Neoheterospilus was also found deeply nested within a large Heterospilus clade, suggesting the paraphyly of the latter genus.

Utility of the DNA barcoding gene fragment for parasitic wasp phylogeny (Hymenoptera: Ichneumonoidea): data release and new measure of taxonomic congruence

The enormous cytochrome oxidase subunit I (COI) sequence database being assembled from the various DNA barcoding projects as well as from independent phylogenetic studies constitutes an almost unprecedented amount of data for molecular systematics, in addition to its role in species identification and discovery. As part of a study of the potential of this gene fragment to improve the accuracy of phylogenetic reconstructions, and in particular, exploring the effects of dense taxon sampling, we have assembled a data set for the hyperdiverse, cosmopolitan parasitic wasp superfamily Ichneumonoidea, including the release of 1793 unpublished sequences. Of approximately 84 currently recognized Ichneumonoidea subfamilies, 2500 genera and 41 000 described species, barcoding 5′‐COI data were assembled for 4168 putative species‐level terminals (many undescribed), representing 671 genera and all but ten of the currently recognized subfamilies. After the removal of identical and near‐identical sequences, the 4174 initial sequences were reduced to 3278. We show that when subjected to phylogenetic analysis using both maximum likelihood and parsimony, there is a broad correlation between taxonomic congruence and number of included sequences. We additionally present a new measure of taxonomic congruence based upon the Simpson diversity index, the Simpson dominance index, which gives greater weight to morphologically recognized taxonomic groups (subfamilies) recovered with most representatives in one or a few contiguous groups or subclusters.

Phylogeny and Niche Conservatism in North and Central American Triatomine Bugs (Hemiptera: Reduviidae: Triatominae), Vectors of Chagas' Disease

The niche conservatism hypothesis states that related species diverge in niche characteristics at lower rates than expected, given their lineage divergence. Here we analyze whether niche conservatism is a common pattern among vector species (Hemiptera: Reduviidae: Triatominae) of Trypanosoma cruzi that inhabit North and Central America, a highly heterogeneous landmass in terms of environmental gradients. Mitochondrial and nuclear loci were used in a multi-locus phylogenetic framework to reconstruct phylogenetic relationships among species and estimate time of divergence of selected clades to draw biogeographic inferences. Then, we estimated similarity between the ecological niche of sister species and tested the niche conservatism hypothesis using our best estimate of phylogeny. Triatoma is not monophyletic. A primary clade with all North and Central American (NCA) triatomine species from the genera Triatoma, Dipetalogaster, and Panstrongylus, was consistently recovered. Nearctic species within the NCA clade (T. p. protracta, T. r. rubida) diverged during the Pliocene, whereas the Neotropical species (T. phyllosoma, T. longipennis, T. dimidiata complex) are estimated to have diverged more recently, during the Pleistocene. The hypothesis of niche conservatism could not be rejected for any of six sister species pairs. Niche similarity between sister species best fits a retention model. While this framework is used here to infer niche evolution, it has a direct impact on spatial vector dynamics driven by human population movements, expansion of transportation networks and climate change scenarios.

Suspended mummies in Aleiodes species (Hymenoptera: Braconidae: Rogadinae) with descriptions of six new species from western Uganda based largely on DNA sequence data

A group of species of the rogadine braconid genus Aleiodes are shown to produce a distinctive mummy, which is “J”‐shaped and is formed after the host larva, in all cases an ennomine geometrid moth, has dropped from a plant suspended in midair by a silk thread. The group includes one described species, A. buzurae He & Chen from China, and a species complex from tropical Africa (W. Uganda). All the African specimens reared from suspended mummies looked morphologically virtually indistinguishable, though there was considerable colour variation that segregated the specimens into five groups. Three gene fragments (nuclear 28S D2‐3 rDNA, the nuclear ITS2 region and part of the mitochondrial cytochrome oxidase 1 gene (CO1)) were sequenced to assess if these specimens represented a single variable species or a complex of morphologically cryptic species. Results show variation in all three gene fragments, with strong signal in the CO1 gene, parsimony analysis of which revealed six well supported groups corresponding to the colour variants, except that two specimens with nearly identical colour differed considerably in their CO1 sequences. Large, and difficult to align, variation was found in the ITS2 fragments, which by eye also supported the same six groupings. Limited variation was found in the 28S fragment, but one position supported monophyly of the two specimens belonging to one of the species circumscribed by the other genes. These groups are considered to correspond to separate species, which are described as new: A. barnardae Quicke & Shaw, A. basutai Quicke & Shaw, A. kanyawarensis Quicke & Shaw, A. kasenenei Quicke & Shaw, A. mubfsi Quicke & Shaw and A. trevelyanae Quicke & Shaw. The possible function of the specialised mummification behaviour is discussed and some observations on rates of hyperparasitism are presented.

Systematics and evolution of gall formation in the plant‐associated genera of the wasp subfamily Doryctinae (Hymenoptera: Braconidae)

Gall formation is a specialised form of phytophagy that consists of abnormal growth of host plant tissue induced by other organisms, principally insects and mites. In the mainly parasitoid wasp subfamily Doryctinae, gall association, represented by gall inducers, inquilines and their parasitoids, is known for species of seven genera. Previous molecular studies recovered few species of six of these genera as monophyletic despite their disparate morphologies. Here, we reconstructed the evolutionary relationships among 47 species belonging to six gall‐associated doryctine genera based on two mitochondrial and two nuclear gene markers. Most of the Bayesian analyses, performed with different levels of incomplete taxa and characters, supported the monophyly of gall‐associated doryctines, with Heterospilus (Heterospilini) as sister group. Percnobracon Kieffer and Jörgensen and Monitoriella Hedqvist were consistently recovered as monophyletic, and the validity of the monotypic Mononeuron was confirmed with respect to Allorhogas Gahan. A nonmonophyletic Allorhogas was recovered, although without significant support. The relationships obtained and the gathered morphological and biological information led us to erect three new genera originally assigned to Psenobolus: Ficobolus gen.n. (F. paniaguai sp.n. and F. jaliscoi sp.n.), Plesiopsenobolus gen.n. (Pl. mesoamericanus sp.n., Pl. plesiomorphus van Achterberg and Marsh comb.n., and Pl. tico sp.n.), and Sabinita gen.n. (S. mexicana sp.n.). The origin of the gall‐associated doryctine clade was estimated to have occurred during the middle Miocene to early Oligocene, 16.33–30.55 Ma. Our results support the origin of true gall induction in the Doryctinae from parasitoidism of other gall‐forming insects. Moreover, adaptations to attack different gall‐forming taxa on various unrelated plant families probably triggered species diversification in the main Allorhogas clade and may also have promoted the independent origin of gall formation on at least three plant groups. Species diversification in the remaining doryctine taxa was probably a result of host shifts within a particular plant taxon and shifts to different plant organs.