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      A revision of the Aleiodes bakeri (Brues) species subgroup of the A. seriatus species group with the descriptions of 18 new species from the Neotropical Region

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          The Aleiodes bakeri (Brues) species subgroup of the A. seriatus species group is defined based on two previously described species, A. bakeri and A. nigristemmaticum (Enderlein), and is greatly expanded in this paper with an identification key, descriptions, and illustrations of 18 new species from the Neotropical Region: A. andinus Shaw & Shimbori, sp. nov.; angustus Shimbori & Shaw, sp. nov.; asenjoi Shimbori & Shaw, sp. nov.; bahiensis Shimbori & Shaw, sp. nov.; barrosi Shimbori & Shaw, sp. nov.; brevicarina Shimbori & Shaw, sp. nov.; coariensis Shimbori & Shaw, sp. nov.; goiasensis Shimbori & Shaw, sp. nov.; gonodontivorus Shaw & Shimbori, sp. nov.; hyalinus Shimbori & Shaw, sp. nov.; inga Shimbori & Shaw, sp. nov.; joaquimi Shimbori & Shaw, sp. nov.; lidiae Shimbori & Shaw, sp. nov.; mabelae Shimbori & Shaw, sp. nov.; maculosus Shimbori & Shaw, sp. nov.; ovatus Shimbori & Shaw, sp. nov.; santarosensis Shaw & Shimbori, sp. nov.; and taurus Shimbori & Penteado-Dias, sp. nov. It is hypothesized that the A. bakeri species subgroup is a monophyletic lineage within the larger and probably artificial A. seriatus species group (those Aleiodes with a comb of flat setae at the apex of the hind tibia), and can be distinguished from other members of the seriatus group by having the hind wing vein r present, although weakly indicated; the hind wing marginal cell suddenly widened at junction of veins RS and r; the subbasal cell of the fore wing mostly glabrous but often with two rows of short setae subapically; glabrous regions of the wings also commonly found in the first subdiscal, discal, and basal cells of the fore wing, and the basal cell of hind wing; ocelli quite large, with the width of a lateral ocellus being distinctly larger than the ocellar-ocular distance; and being relatively large Aleiodes species with body almost entirely brownish yellow or reddish brown. In addition, a new replacement name, Aleiodes buntikae Shimbori & Shaw, nom. nov., is proposed for the species formerly called Aleiodes ( Hemigyroneuron) bakeri Butcher & Quicke, 2011.

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          Extreme diversity of tropical parasitoid wasps exposed by iterative integration of natural history, DNA barcoding, morphology, and collections.

          We DNA barcoded 2,597 parasitoid wasps belonging to 6 microgastrine braconid genera reared from parapatric tropical dry forest, cloud forest, and rain forest in Area de Conservación Guanacaste (ACG) in northwestern Costa Rica and combined these data with records of caterpillar hosts and morphological analyses. We asked whether barcoding and morphology discover the same provisional species and whether the biological entities revealed by our analysis are congruent with wasp host specificity. Morphological analysis revealed 171 provisional species, but barcoding exposed an additional 142 provisional species; 95% of the total is likely to be undescribed. These 313 provisional species are extraordinarily host specific; more than 90% attack only 1 or 2 species of caterpillars out of more than 3,500 species sampled. The most extreme case of overlooked diversity is the morphospecies Apanteles leucostigmus. This minute black wasp with a distinctive white wing stigma was thought to parasitize 32 species of ACG hesperiid caterpillars, but barcoding revealed 36 provisional species, each attacking one or a very few closely related species of caterpillars. When host records and/or within-ACG distributions suggested that DNA barcoding had missed a species-pair, or when provisional species were separated only by slight differences in their barcodes, we examined nuclear sequences to test hypotheses of presumptive species boundaries and to further probe host specificity. Our iterative process of combining morphological analysis, ecology, and DNA barcoding and reiteratively using specimens maintained in permanent collections has resulted in a much more fine-scaled understanding of parasitoid diversity and host specificity than any one of these elements could have produced on its own.
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            DNA barcodes affirm that 16 species of apparently generalist tropical parasitoid flies (Diptera, Tachinidae) are not all generalists.

            Many species of tachinid flies are viewed as generalist parasitoids because what is apparently a single species of fly has been reared from many species of caterpillars. However, an ongoing inventory of the tachinid flies parasitizing thousands of species of caterpillars in Area de Conservación Guanacaste, northwestern Costa Rica, has encountered >400 species of specialist tachinids with only a few generalists. We DNA-barcoded 2,134 flies belonging to what appeared to be the 16 most generalist of the reared tachinid morphospecies and encountered 73 mitochondrial lineages separated by an average of 4% sequence divergence. These lineages are supported by collateral ecological information and, where tested, by independent nuclear markers (28S and ITS1), and we therefore view these lineages as provisional species. Each of the 16 apparently generalist species dissolved into one of four patterns: (i) a single generalist species, (ii) a pair of morphologically cryptic generalist species, (iii) a complex of specialist species plus a generalist, or (iv) a complex of specialists with no remaining generalist. In sum, there remained 9 generalist species among the 73 mitochondrial lineages we analyzed, demonstrating that a generalist lifestyle is possible for a tropical caterpillar parasitoid fly. These results reinforce the emerging suspicion that estimates of global species richness are likely underestimates for parasitoids (which may constitute as much as 20% of all animal life) and that the strategy of being a tropical generalist parasitic fly may be yet more unusual than has been envisioned for tachinids.
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              Skeletal Morphology of Opius dissitus and Biosteres carbonarius (Hymenoptera: Braconidae), with a Discussion of Terminology

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

                Author and article information

                Pensoft Publishers
                27 August 2020
                : 964
                : 41-107
                [1 ] Department of Ecosystem Science and Management, University of Wyoming, Laramie, Wyoming 82071-3354, USA University of Wyoming Laramie United States of America
                [2 ] ESALQ/USP, Departamento de Entomologia e Acarologia – LEA, Avenida Pádua Dias, 11 Piracicaba/SP, CEP 13418-900, Brazil ESALQ/USP, Departamento de Entomologia e Acarologia – LEA Piracicaba Brazil
                [3 ] Universidade Federal de São Carlos, Rodovia Washington Luiz, km 235, CEP 13 565-905, São Carlos, SP, Brazil Universidade Federal de São Carlos São Carlos Brazil
                Author notes
                Corresponding author: Eduardo M. Shimbori ( shimbori@ 123456usp.br )

                Academic editor: Kees van Achterberg

                Scott R. Shaw, Eduardo M. Shimbori, Angelica M. Penteado-Dias

                This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

                National Science Foundation grants DEB-10-20751 and DEB 14-42110 (Dimensions of Biodiversity Program) McIntire-Stennis Grant Project number WYO-530-14 INCT-HYMPAR (Instituto Nacional de Ciência e Tecnologia dos Hymenoptera Parasitoides) USDA National Institute of Food and Agriculture, McIntire-Stennis project 1021111 Coordenação de Aperfeiçoamento de Pessoal de Nível Superior – Brasil (CAPES) – Finance Code 001
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