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Declines in biodiversity generated by anthropogenic stressors at both species and population levels can alter emergent processes instrumental to ecosystem function and resilience. As such, understanding the role of biodiversity in ecosystem function and its response to climate perturbation is increasingly important, especially in tropical systems where responses to changes in biodiversity are less predictable and more challenging to assess experimentally. Using large scale transplant experiments conducted at five neotropical sites, we documented the impacts of changes in intraspecific and interspecific plant richness in the genusPiperon insect herbivory, insect richness, and ecosystem resilience to perturbations in water availability. We found that reductions of both intraspecific and interspecificPiperdiversity had dramatic and site specific effects on herbivory, herbivorous insect richness, and plant mortality. Ecosystem responses to reduced intraspecific richness were often similar in magnitude to responses to reduced interspecific richness. Increased water availability reduced herbivory by 4.2% overall, and the response of herbivorous insect richness and herbivory to water availability was altered by both intra and interspecific richness in a site contingent manner. Our results underscore the role of intraspecific and interspecific richness as foundations of ecosystem function, and the importance of community specific contingencies in controlling function in complex tropical systems.

Population divergence can occur due to mechanisms associated with geographic isolation and/or due to selection associated with different ecological niches. Much of the evidence for selection-driven speciation has come from studies of specialist insect herbivores that use different host plant species; however, the influence of host plant use on population divergence of generalist herbivores remains poorly understood. We tested how diet breadth, host plant species and geographic distance influence population divergence of the fall webworm (Hyphantria cunea; FW). FW is a broadly distributed, extreme generalist herbivore consisting of two morphotypes that have been argued to represent two different species: black-headed and red-headed. We characterized the differentiation of FW populations at two geographic scales. We first analysed the influence of host plant and geographic distance on genetic divergence across a broad continental scale for both colour types. We further analysed the influence of host plant, diet breadth and geographic distance on divergence at a finer geographic scale focusing on red-headed FW in Colorado. We found clear genetic and morphological distinction between red- and black-headed FW, and Colorado FW formed a genetic cluster distinct from other locations. Although both geographic distance and host plant use were correlated with genetic distance, geographic distance accounted for up to 3× more variation in genetic distance than did host plant use. As a rare study investigating the genetic structure of a widespread generalist herbivore over a broad geographic range (up to 3,000 km), our study supports a strong role for geographic isolation in divergence in this system.

Pelamera atra (Rondani, 1861) is an enigmatic species of bristle fly that has been challenging dipterologists with regard to its taxonomic position within the family Tachinidae since it was first described. The species is rarely collected and is only known from a handful of female specimens. The first ever male specimen of P. atra was collected in southern Italy in 2017, and it allowed for the discovery of its unique terminalia. The present paper provides the first description of a male of P. atra and presents new insights about its phylogenetic position within the family based on molecular data. A maximum likelihood analysis was conducted on a selection of tachinid taxa using sequences of two nuclear protein-coding genes (CAD and MCS) to formulate the first hypothesis concerning the phylogenetic placement of P. atra. Phylogenetic analyses reconstructed P. atra as a member of the Tachininae sister to the (Loewiini + (Polideini + Ernestiini)) clade. This reconstruction challenges previous attempts at classifying Pelamera based on female external morphology, which instead suggested a close affinity to the tribes Brachymerini or Minthoini. Due to the morphological distinctiveness of this taxon, we here propose the placement of P. atra in the monotypic tribe Pelamerini stat. rev.

The family Tachinidae (“bristle flies”) is the most diverse and ecologically important group of insect parasitoids outside the parasitic wasps. It is among the most species rich families of flies (Diptera) and has experienced a recent adaptive radiation across the globe. We make use of a molecular phylogeny of the family to examine its rapid radiation and explore the traits of tachinid lineages that may have contributed to variation in their diversification. We apply a range of diversification analyses to assess the consistency and robustness of effects. We find that the Tachinidae are among the most rapidly diversifying families of animals. Six to eight clades of bristle flies, distributed across the phylogeny, exhibit strong evidence of accelerated diversification. Our results suggest that the use of holometabolous insect larvae, and specifically caterpillars (Lepidoptera), as hosts, is associated with increased diversification rates. However, these effects were inconsistent across analyses. We detected little influence of oviposition strategy (egg type) or host feeding habit, and we recovered evidence that unmeasured “hidden” traits may explain greater variance in diversification. We evaluated the strengths and weaknesses of different Maximum Likelihood and Bayesian approaches for analysing diversification and the potential for extrinsic factors, such as geography, to influence patterns of richness and diversification. In general, we conclude that although certain traits may provide opportunities for diversification, whether this is capitalized on may depend on additional traits and/or historical contingency.

Figs and their associated mutualistic and parasitic wasps have been a focus of intensive ecological and evolutionary research due to their diversity, unusual reproductive biology, and highly coevolved interspecific relationships. Due to the ecological dependence of their interactions, fig wasps were once considered to be fig-species specific and to cospeciate with their hosts, however, a growing body of evidence reveals mixed support for species specificity and the importance of additional evolutionary processes (e.g., host switching) structuring these long-term interactions. Our research on the genus Idarnes Walker, 1843 (Hymenoptera, Agaonidae), a common non-pollinating wasp of New World fig flowers, reveals a community in which multiple wasp species coexist on the same host in space and time. Using both molecular and morphological data, we identify five distinct Idarnes lineages associated with a single host fig species, Ficus petiolaris Kunth, 1817 (Rosales, Moraceae). A comprehensive phylogenetic analysis including Idarnes species from numerous host fig species reveals that the lineages associated with F. petiolaris do not form a monophyletic group but are distantly related, suggesting multiple independent colonization events and subsequent diversification. Morphological and ecological data provide support that the wasps are partitioning niches within the figs, explaining the coexistence of these diverse lineages on the same host fig. These results, coupled with a growing body of research on pollinating and non-pollinating fig wasps, bring into focus a more dynamic picture of fig and fig wasp coevolution and highlight how wasp lineage divergence and niche partitioning contributes to increased species diversity and community structure on a single fig host.

A multi-trophic perspective improves understanding of the ecological and evolutionary consequences of rapid environmental change on insect herbivores. Loss of specialized enemies due to human impacts is predicted to dramatically reduce the number of tritrophic niches of herbivores compared to a bitrophic niche perspective. Habitat fragmentation and climate change promote the loss of both specialist enemies and herbivores, favoring ecological generalism across trophic levels. Species invasion can fundamentally alter trophic interactions toward various outcomes and contributes to ecological homogenization. Adaptive evolution on ecological timescales is expected to dampen tritrophic instabilities and diversify niches, yet its ability to compensate for tritrophic niche losses in the short term is unclear.

Associated data is available at: https://doi.org/10.5061/dryad.595718c; International audience; Habitat destruction is the single greatest anthropogenic threat to biodiversity. Decades of research on this issue have led to the accumulation of hundreds of data sets comparing species assemblages in larger, intact, habitats to smaller, more fragmented, habitats. Despite this, little synthesis or consensus has been achieved, primarily because of non-standardized sampling methodology and analyses of notoriously scale-dependent response variables (i.e., species richness). To be able to compare and contrast the results of habitat fragmentation on species’ assemblages, it is necessary to have the underlying data on species abundances and sampling intensity, so that standardization can be achieved. To accomplish this, we systematically searched the literature for studies where abundances of species in assemblages (of any taxa) were sampled from many habitat patches that varied in size. From these, we extracted data from several studies, and contacted authors of studies where appropriate data were collected but not published, giving us 117 studies that compared species assemblages among habitat fragments that varied in area. Less than one-half (41) of studies came from tropical forests of Central and South America, but there were many studies from temperate forests and grasslands from all continents except Antarctica. Fifty-four of the studies were on invertebrates (mostly insects), but there were several studies on plants (15), birds (16), mammals (19), and reptiles and amphibians (13). We also collected qualitative information on the length of time since fragmentation. With data on total and relative abundances (and identities) of species, sampling effort, and affiliated meta-data about the study sites, these data can be used to more definitively test hypotheses about the role of habitat fragmentation in altering patterns of biodiversity. There are no copyright restrictions. Please cite this data paper and the associated Dryad data set if the data are used in publications.

Article, The Glaurocarini are a small Old World tribe of tachinids belonging to the subfamily Tachininae. Two genera are currently recognized, Glaurocara Thomson with 16 species and Semisuturia Malloch with eight species. In this study we describe Semisuturia moffattensis Inclán, O’Hara, Stireman & Cerretti sp. n. from Queensland and New South Wales and compare it with congeners as well as other glaurocarines. The new species is readily identifiable among world glaurocarines by having a row of setae on lower 2/3 of facial ridge. We further evaluate the monophyly of the Glaurocarini on the basis of morphological characters of both adult and larval stages. A molecular phylogenetic analysis also supports monophyly of the tribe but does not support a close relationship between Glaurocarini and Ormiini as has been suggested previously. Finally, we provide new morphological evidence from both adults and first instar larvae to support the monophyly of both Semisuturia and Glaurocara. The Glaurocarini are a small Old World tribe of tachinids belonging to the subfamily Tachininae. Two genera are currently recognized, Glaurocara Thomson with 16 species and Semisuturia Malloch with eight species. In this study we describe Semisuturia moffattensis Inclán, O’Hara, Stireman & Cerretti sp. n. from Queensland and New South Wales and compare it with congeners as well as other glaurocarines. The new species is readily identifiable among world glaurocarines by having a row of setae on lower 2/3 of facial ridge. We further evaluate the monophyly of the Glaurocarini on the basis of morphological characters of both adult and larval stages. A molecular phylogenetic analysis also supports monophyly of the tribe but does not support a close relationship between Glaurocarini and Ormiini as has been suggested previously. Finally, we provide new morphological evidence from both adults and first instar larvae to support the monophyly of both Semisuturia and Glaurocara., final draft, peerReviewed

Extensive rearing of monarch larvae (Danaus plexippus L.) through the citizen science Monarch Larva Monitoring Project (MLMP) revealed that monarchs' primary parasitoids are flies in the family Tachinidae and that these parasitoids result in appreciable larval mortality. We document the tachinid community that attacks monarchs in the United States, evaluate their relative frequency, and examine variation in their specificity, oviposition strategy, and use of host stages. Based on results of rearing >20,000 monarchs by MLMP volunteers, overall parasitism by tachinids across life stages was 9.8% (17% for monarchs collected as fifth instars). We identified the flies that emerged from 466 monarch hosts, and found seven Tachinidae species. In decreasing order of frequency, these included Lespesia archippivora (Riley), Hyphantrophaga virilis (Aldrich & Webber), Compsilura concinnata (Meigen), Leschenaultia n. sp., Madremyia saundersii (Williston), Lespesia sp., and Nilea erecta (Coquillett). Lespesia sp., Leschenaultia n. sp., and N. erecta had not been previously reported as monarch parasitoids, and Leschenaultia n. sp. is apparently undescribed. We include new state records (Texas and Iowa) for C. concinnata. Lespesia archippivora and C. concinnata were overrepresented as parasitoids of later instars and were absent from monarchs collected as eggs, but H. virilis and Leschenaultia sp., which lay their eggs on foliage, were reared from caterpillars collected as eggs. To our knowledge, we include the first report of multiparasitism of monarchs, in which more than one parasitoid species emerged from a host. The biology of the tachinid parasitoids we identified and their relationship with monarchs is examined.

Most insect species remain unknown and for most described species, we know nothing about their ecology and interactions. Understanding the structure of species interaction webs can provide insight into geographic patterns and ecological drivers of diversity, the stability of communities, and energy flow through ecological systems. We analysed a tachinid parasitoid-caterpillar food web from the Ecuadorian Andes based on >70 000 caterpillar collections. Our goals were to assess how host taxon and feeding niche affects parasitism by flies and wasps, evaluate perceptions of broad diet breadth in tachinids, characterise the structure of the food web, and assess how sample size affects inferences about community structure. We found high levels of parasitism (>28%) by tachinid and wasp parasitoids and a rich tachinid community (279 spp.). Concealed caterpillars experienced low parasitism by tachinids and were more likely to be attacked by species with indirect oviposition. Indirect ovipositing tachinids had greater host ranges, but this is confounded with tachinid phylogeny. Caterpillar families exhibited nearly opposite patterns of susceptibility to parasitism by tachinids and wasps. Network metrics indicate that this tachinid-host community is relatively compartmentalised and nested, and though some taxa display broad host ranges, the network as a whole is moderately specialised due to constrained host ranges of many tachinid species. This study establishes an exemplar tropical tachinid-host food web, providing insight into the diversity and interactions of these important parasitoids. Additional studies are needed to determine how these interactions vary across environmental and geographic gradients.

Little is known about the biology or conservation status of Lucanus elaphus Fabricius in North America despite well-documented declines of a related species, Lucanus cervus (L.), in Europe. This study provides information critical to developing conservation plans for L. elaphus including the species’ larval substrate requirements, genetic data and range-wide estimates of habitat suitability. In Mississippi floodplain forests, larval L. elaphus were recovered from a wide range of log sizes and rot types and were either found tunnelling within the wood or feeding beneath logs at the soil–wood interface. The species appears to require 1–2 years to complete development, exhibits a 1:1 sex ratio and is parasitised by Zelia vertebrata (Say) (Diptera: Tachinidae). Flight intercept traps placed at three heights at both the edge and interior of hardwood-dominated forests in Georgia yielded six adult male L. elaphus, all of which were captured in traps placed at 15 m on the forest edge. Because L. elaphus larvae are morphologically indistinguishable from related species, DNA sequence data from the mitochondrial cytochrome oxidase I gene were generated to facilitate molecular identification. Genetic data revealed modest intra-specific variation, with up to 1.3% sequence divergence among haplotypes sampled from the same forest. Based on assembled occurrence records, ecological niche models suggest that environmental conditions are suitable for L. elaphus across much of the southeastern United States, provided that adequate lowland forest cover and dead wood substrates are available.

Gall midges (Cecidomyiidae) constitute one of the largest and most diverse families of Diptera, with close to 6600 described species and thousands of undescribed species worldwide. The family is divided into six subfamilies, the five basal ones comprising only fungivorous taxa, whereas the largest, youngest and most diverse subfamily Cecidomyiinae includes fungivorous as well as herbivorous and predatory species. The currently accepted classification of the Cecidomyiinae is morphology-based, and the few phylogenetic inferences that have previously been suggested for it were based on fragmentary or limited datasets. In a first comprehensive phylogenetic analysis of the Cecidomyiinae we sampled 142 species representing 88 genera of 13 tribes from all feeding guilds and zoogeographic regions in order to test the validity of the systematic division of the subfamily and gain insight into patterns of diversification and the evolution of feeding modes. We used sequences from five mitochondrial and nuclear genes to reconstruct maximum likelihood and Bayesian, time-calibrated phylogenies and conducted ancestral state reconstruction of feeding modes. Our results corroborate to a great extent the morphology-based classification of the Cecidomyiinae, with strong support for all supertribes and tribes, all were apparently established in the Upper Cretaceous concordant with the major radiation of angiosperms. We infer that transitions from fungus-feeding to plant-feeding occurred only once or twice in the evolution of the subfamily and that predation evolved only once, contrary to previous hypotheses. All herbivorous clades in the subfamily are very species rich and have diversified at a significantly greater rate than expected, but we found no support for the assertion that herbivorous clades associated with symbiotic fungi in their galls diversify faster than clades that do not have such associations. Currently available data also do not support the hypothesis that symbiotic clades have broader host ranges than non-symbiotic clades.

Family Polleniidae Brauer & Bergenstamm, 1889 stat. rev. Diagnosis. Small to medium-sized oestroid flies varying from yellow to black in ground colour. Facial sclerite at least weakly carinate [with few exceptions, e.g. Pollenia griseotomentosa (Jacentkovsk��)]. Stem vein bare dorsally. Anal vein not reaching wing margin. Posterodorsal margin of hind coxa bare. Prosternum and proepisternal depression bare. Postalar wall setose (occasionally bare in small specimens of Morinia). Female: ovipositor sclerite length moderate; sternite 8 of ovipositor elongate with apex entire; cerci long and narrow. Male: ventral and ventrolateral surface of distalmost parts of distiphallus smooth., Published as part of Cerretti, Pierfilippo, Stireman III, John O., Badano, Davide, Gisondi, Silvia, Rognes, Knut, Giudice, Giuseppe Lo & Pape, Thomas, 2019, Reclustering the cluster flies (Diptera: Oestroidea, Polleniidae), pp. 957-972 in Systematic Entomology 44 on page 964, DOI: 10.1111/syen.12369, http://zenodo.org/record/3402822, {"references":["Brauer, F. & von Bergenstamm, J. E. (1889) Die Zweiflugler des kaiserlichen Museums zu Wien. IV. Vorarbeiten zu einer Monographie der Muscariae Schizometopa (exclusive Anthomyidae). Pars I, Vol. 56, pp. 69 - 180. Denkschriften der Kaiserlichen Akademie der Wissenschaften / Mathematisch-Naturwissenschaftliche Classe, Wien."]}

We reconstructed phylogenetic relationships within the diverse parasitoid fly family Tachinidae using four nuclear loci (7800 bp) and including an exceptionally large sample of more than 500 taxa from around the world. The position of the earthworm-parasitizing Polleniinae (Calliphoridae s.l.) as sister to Tachinidae is strongly supported. Our analyses recovered each of the four tachinid subfamilies and most recognized tribes, with some important exceptions in the Dexiinae and Tachininae. Most notably, the tachinine tribes Macquartiini and Myiophasiini form a clade sister to all other Tachinidae, and a clade of Palpostomatini is reconstructed as sister to Dexiinae + Phasiinae. Although most nodes are well-supported, relationships within several lineages that appear to have undergone rapid episodes of diversification (basal Dexiinae and Tachininae, Blondeliini) were poorly resolved. Reconstructions of host use evolution are equivocal, but generally support the hypothesis that the ancestral host of tachinids was a beetle and that subsequent host shifts to caterpillars may coincide with accelerated diversification. Evolutionary reconstructions of reproductive strategy using alternative methods were incongruent, however it is most likely that ancestral tachinids possessed unincubated, thick shelled eggs from which incubated eggs evolved repeatedly, potentially expanding available host niches. These results provide a broad foundation for understanding the phylogeny and evolution of this important family of parasitoid insects. We hope it will serve as a framework to be used in concert with morphology and other sources of evidence to revise the higher taxonomic classification of Tachinidae and further explore their evolutionary history and diversification.

Most studies of adaptive radiation in animals focus on resource competition as the primary driver of trait divergence. The roles of other ecological interactions in shaping divergent phenotypes during such radiations have received less attention. We evaluate natural enemies as primary agents of diversifying selection on the phenotypes of an actively diverging lineage of gall midges on tall goldenrod. In this system, the gall of the midge consists of a biotrophic fungal symbiont that develops on host-plant leaves and forms distinctly variable protective carapaces over midge larvae. Through field studies, we show that fungal gall morphology, which is induced by midges (i.e., it is an extended phenotype), is under directional and diversifying selection by parasitoid enemies. Overall, natural enemies disruptively select for either small or large galls, mainly along the axis of gall thickness. These results imply that predators are driving the evolution of phenotypic diversity in symbiotic defense traits in this system and that divergence in defensive morphology may provide ecological opportunities that help to fuel the adaptive radiation of this genus of midges on goldenrods. This enemy-driven phenotypic divergence in a diversifying lineage illustrates the potential importance of consumer-resource and symbiotic species interactions in adaptive radiation.

Study of all flies (Diptera) collected for one year from a four-hectare (150 x 266 meter) patch of cloud forest at 1,600 meters above sea level at Zurquí de Moravia, San José Province, Costa Rica (hereafter referred to as Zurquí), revealed an astounding 4,332 species. This amounts to more than half the number of named species of flies for all of Central America. Specimens were collected with two Malaise traps running continuously and with a wide array of supplementary collecting methods for three days of each month. All morphospecies from all 73 families recorded were fully curated by technicians before submission to an international team of 59 taxonomic experts for identification. Overall, a Malaise trap on the forest edge captured 1,988 species or 51% of all collected dipteran taxa (other than of Phoridae, subsampled only from this and one other Malaise trap). A Malaise trap in the forest sampled 906 species. Of other sampling methods, the combination of four other Malaise traps and an intercept trap, aerial/hand collecting, 10 emergence traps, and four CDC light traps added the greatest number of species to our inventory. This complement of sampling methods was an effective combination for retrieving substantial numbers of species of Diptera. Comparison of select sampling methods (considering 3,487 species of non-phorid Diptera) provided further details regarding how many species were sampled by various methods. Comparison of species numbers from each of two permanent Malaise traps from Zurquí with those of single Malaise traps at each of Tapantí and Las Alturas, 40 and 180 km distant from Zurquí respectively, suggested significant species turnover. Comparison of the greater number of species collected in all traps from Zurquí did not markedly change the degree of similarity between the three sites, although the actual number of species shared did increase. Comparisons of the total number of named and unnamed species of Diptera from four hectares at Zurquí is equivalent to 51% of all flies named from Central America, greater than all the named fly fauna of Colombia, equivalent to 14% of named Neotropical species and equal to about 2.7% of all named Diptera worldwide. Clearly the number of species of Diptera in tropical regions has been severely underestimated and the actual number may surpass the number of species of Coleoptera. Various published extrapolations from limited data to estimate total numbers of species of larger taxonomic categories (e.g., Hexapoda, Arthropoda, Eukaryota, etc.) are highly questionable, and certainly will remain uncertain until we have more exhaustive surveys of all and diverse taxa (like Diptera) from multiple tropical sites. Morphological characterization of species in inventories provides identifications placed in the context of taxonomy, phylogeny, form, and ecology. DNA barcoding species is a valuable tool to estimate species numbers but used alone fails to provide a broader context for the species identified.

Made available in DSpace on 2019-10-06T15:56:52Z (GMT). No. of bitstreams: 0 Previous issue date: 2018-12-01 Estimations of tropical insect diversity generally suffer from lack of known groups or faunas against which extrapolations can be made, and have seriously underestimated the diversity of some taxa. Here we report the intensive inventory of a four-hectare tropical cloud forest in Costa Rica for one year, which yielded 4332 species of Diptera, providing the first verifiable basis for diversity of a major group of insects at a single site in the tropics. In total 73 families were present, all of which were studied to the species level, providing potentially complete coverage of all families of the order likely to be present at the site. Even so, extrapolations based on our data indicate that with further sampling, the actual total for the site could be closer to 8000 species. Efforts to completely sample a site, although resource-intensive and time-consuming, are needed to better ground estimations of world biodiversity based on limited sampling. Entomology Section Natural History Museum of Los Angeles County, 900 Exposition Boulevard Royal British Columbia Museum and the American Museum of Natural History, 691-8th Ave. SE Department of Plant and Environmental Sciences Poole Agricultural Center Clemson University, 130 McGinty Court, E-143 Depto. de Biologia FFCLRP Universidade de São Paulo, Av. Bandeirantes 3900 Great Lakes Forestry Centre Canadian Forest Service Natural Resources Canada, 1219 Queen St. E. Australian Museum, 1 William Street Department of Natural Resource Sciences McGill University Macdonald Campus Canadian National Collection of Insects Invertebrate Biodiversity Agriculture and Agri-Food Canada, K.W. Neatby Building, 960 Carling Avenue Department of Biological Sciences Spaulding Hall University of New Hampshire Department of Biological Sciences Wright State University, 3640 Colonel Glenn Hwy Instituto Federal do Triângulo Mineiro – Campus Uberaba. Rua João Batista Ribeiro 4000 Distrito Industrial II Departamento de Zoologia Universidade Federal do Paraná Jardim das Américas Mississippi Entomological Museum Mississippi State University, 100 Old Highway 12, P.O. Drawer 9775 Department of Biology Western Kentucky University Integrative Research Center Field Museum of Natural History Independent Investigator California State Collection of Arthropods, 2683 Tam O’ Shanter Dr., El Dorado Hills California Department of Food and Agriculture California State Collection of Arthropods, 3294 Meadowview Rd. The Academy of Natural Sciences of Drexel University, 1900 Ben Franklin Parkway American Museum of Natural History, Central Park West at 79th St. Department of Entomology University of California Riverside, 900 University Ave. Zoological Museum Biodiversity Unit FI-20014 University of Turku Instituto de Ecología A.C. (INECOL) Red Ambiente y Sustentabilidad, Carretera Antigua a Coatepec 351, Col El Haya, Xalapa Station Linné, Ölands Skogsby 161 I. I. Schmalhausen Institute of Zoology of the National Academy of Sciences of Ukraine, Bogdan Chmielnicki St. 15 Universidad de Panama Department of Natural History University Museum of Bergen University of Bergen, P.O. Box 7800 Agriculture and Agri-Food Canada, 960 Carling Avenue School of Environmental Sciences University of Guelph Department of Entomology Smithsonian Institution, PO Box 37012, MRC 169 Natural History Museum of Denmark, Universitetsparken 15 Institute of Evolutionary Biology and Environmental Studies University of Zurich, Winterthurerstrasse 190 Systematic Entomology Laboratory USDA ARS c/o National Museum of Natural History, MRC-168, P.O. Box 37012 Departamento de Sistemática e Ecologia CCEN Universidade Federal da Paraíba, Castelo Branco, s/n Research Institute for Nature and Forest (INBO), Kliniekstraat 25 Research Group Terrestrial Ecology (TEREC) Ghent University, K.L.Ledeganckstraat 35 Entomology Unit Royal Belgian Institute for Natural Sciences (RBINS), Vautierstraat 29 USDA Forest Service Rocky Mountain Research Station Forestry Sciences Laboratory, 1648 S. 7th Avenue Department of Biological Sciences Bishop’s University, 2600 College Street UNESP - Univ Estadual Paulista Faculdade de Ciências Agrárias e Veterinárias Departamento de Morfologia e Fisiologia Animal; Via de Acesso Prof. Paulo Donato Castellane s/n Canadian National Collection of Insects & Canadian Food Inspection Agency OPL-Entomology, K.W. Neatby Bldg., C.E.F., 960 Carling Ave. Department of Biological Sciences University of Calgary, 2500 University Drive NW Finnish Museum of Natural History Zoology Unit University of Helsinki Washington State University, 2533 Inter Avenue Bohart Museum of Entomology University of California, One Shields Avenue Instituto Nacional de Biodiversidad (INBio) UNESP - Univ Estadual Paulista Faculdade de Ciências Agrárias e Veterinárias Departamento de Morfologia e Fisiologia Animal; Via de Acesso Prof. Paulo Donato Castellane s/n

Molecular phylogenetic studies at all taxonomic levels often infer rapid radiation events based on short, poorly resolved internodes. While such rapid episodes of diversification are an important and widespread evolutionary phenomenon, much of this poor phylogenetic resolution may be attributed to the continuing widespread use of “traditional” markers (mitochondrial, ribosomal, and some nuclear protein-coding genes) that are often poorly suited to resolve difficult, higher-level phylogenetic problems. Here we reconstruct phylogenetic relationships among a representative set of taxa of the parasitoid fly family Tachinidae and related outgroups of the superfamily Oestroidea. The Tachinidae are one of the most species rich, yet evolutionarily recent families of Diptera, providing an ideal case study for examining the differential performance of loci in resolving phylogenetic relationships and the benefits of adding more loci to phylogenetic analyses. We assess the phylogenetic utility of nine genes including both traditional genes (e.g., CO1 mtDNA, 28S rDNA) and nuclear protein-coding genes newly developed for phylogenetic analysis. Our phylogenetic findings, based on a limited set of taxa, include: a close relationship between Tachinidae and the calliphorid subfamily Polleninae, monophyly of Tachinidae and the subfamilies Exoristinae and Dexiinae, subfamily groupings of Dexiinae + Phasiinae and Tachininae + Exoristinae, and robust phylogenetic placement of the somewhat enigmatic genera Strongygaster, Euthera, and Ceracia. In contrast to poor resolution and phylogenetic incongruence of “traditional genes,” we find that a more selective set of highly informative genes is able to more precisely identify regions of the phylogeny that experienced rapid radiation of lineages, while more accurately depicting their phylogenetic context. Although much expanded taxon sampling is necessary to effectively assess the monophyly of and relationships among major tachinid lineages and their relatives, we show that a small number of well-chosen nuclear protein-coding genes can successfully resolve even difficult phylogenetic problems.

Calyptrate flies include about 22,000 extant species currently classified into Hippoboscoidea (tsetse, louse, and bat flies), the muscoid grade (house flies and relatives) and the Oestroidea (blow flies, bot flies, flesh flies, and relatives). Calyptrates are abundant in nearly all terrestrial ecosystems, often playing key roles as decomposers, parasites, parasitoids, vectors of pathogens, and pollinators. For oestroids, the most diverse group within calyptrates, definitive fossils have been lacking. The first unambiguous fossil of Oestroidea is described based on a specimen discovered in amber from the Dominican Republic. The specimen was identified through digital dissection by CT scans, which provided morphological data for a cladistic analysis of its phylogenetic position among extant oestroids. The few known calyptrate fossils were used as calibration points for a molecular phylogeny (16S, 28S, CAD) to estimate the timing of major diversification events among the Oestroidea. Results indicate that: (a) the fossil belongs to the family Mesembrinellidae, and it is identified and described as Mesembrinella caenozoica sp. nov.; (b) the mesembrinellids form a sister clade to the Australian endemic Ulurumyia macalpinei (Ulurumyiidae) (McAlpine’s fly), which in turn is sister to all remaining oestroids; (c) the most recent common ancestor of extant Calyptratae lived just before the K–Pg boundary (ca. 70 mya); and (d) the radiation of oestroids began in the Eocene (ca. 50 mya), with the origin of the family Mesembrinellidae dated at ca. 40 mya. These results provide new insight into the timing and rate of oestroid diversification and highlight the rapid radiation of some of the most diverse and ecologically important families of flies. ZooBank accession number–urn:lsid:zoobank.org:pub:0DC5170B-1D16-407A-889E-56EED3FE3627.

Habitat fragmentation and invasive species are two of the most prominent threats to terrestrial ecosystems. Few studies have examined how these factors interact to influence the diversity of natural communities, particularly primary consumers. Here, we examined the effects of forest fragmentation and invasion of exotic honeysuckle (Lonicera maackii, Caprifoliaceae) on the abundance and diversity of the dominant forest herbivores: woody plant-feeding Lepidoptera. We systematically surveyed understory caterpillars along transects in 19 forest fragments over multiple years in southwestern Ohio and evaluated how fragment area, isolation, tree diversity, invasion by honeysuckle and interactions among these factors influence species richness, diversity and abundance. We found strong seasonal variation in caterpillar communities, which responded differently to fragmentation and invasion. Abundance and richness increased with fragment area, but these effects were mitigated by high levels of honeysuckle, tree diversity, landscape forest cover, and large recent changes in area. Honeysuckle infestation was generally associated with decreased caterpillar abundance and diversity, but these effects were strongly dependent on other fragment traits. Effects of honeysuckle on abundance were moderated when fragment area, landscape forest cover and tree diversity were high. In contrast, negative effects of honeysuckle invasion on caterpillar diversity were most pronounced in fragments with high tree diversity and large recent increases in area. Our results illustrate the complex interdependencies of habitat fragmentation, plant diversity and plant invasion in their effects on primary consumers and emphasize the need to consider these processes in concert to understand the consequences of anthropogenic habitat change for biodiversity.

The oestroid family Tachinidae represents one of the most diverse lineages of insect parasitoids. Despite their broad distribution, diversity and important role as biological control agents, the phylogeny of this family remains poorly known. Here, we review the history of tachinid systematics and present the first quantitative phylogenetic analysis of the family based on morphological data. Cladistic analyses were conducted using 135 morphological characters from 492 species belonging to 180 tachinid genera, including the four currently recognized subfamilies (Dexiinae, Exoristinae, Phasiinae, Tachininae) and all major tribes. We used characters of eggs, first-instar larvae and adults of both sexes. We examined the effects of implied weighting by reanalysing the data with varying concavity factors. Our analysis generally supports the subfamily groupings Dexiinae + Phasiinae and Tachininae + Exoristinae, with only the Exoristinae and the Phasiinae reconstructed as monophyletic assemblages under a wide range of weighting schemes. Under these conditions, the Dexiinae, which were previously considered a well-established monophyletic assemblage, are reconstructed as being paraphyletic with respect to the Phasiinae. The Tachininae are reconstructed as a paraphyletic grade from which the monophyletic Exoristinae arose. The Exoristinae are reconstructed as a monophyletic lineage, but phylogenetic relationships within the subfamily are largely unresolved. We further explored the evolution of oviposition strategy and found that the oviparous groups are nested within ovolarviparous assemblages, suggesting that ovipary may have evolved several times independently from ovolarviparous ancestors. This counterintuitive pattern is a novel hypothesis suggested by the results of this analysis. Finally, two major patterns emerge when considering host associations across our phylogeny under equal weights: (i) although more than 60% of tachinids are parasitoids of Lepidoptera larvae, none of the basal clades is unambiguously associated with Lepidoptera as a primitive condition, suggesting that tachinids were slow to colonize these hosts, but then radiated extensively on them; and (ii) there is general agreement between host use and monophyly of the major lineages.

Understanding the evolutionary reasons for patterns of chemical defense in plants is an ongoing theoretical and empirical challenge. The goal is to develop a model that can reliably predict how defenses are distributed within the plant over space and time. This is difficult given that evolutionary, ecological, and physiological processes and tradeoffs can operate over different spatial and temporal scales. We evaluated the major predictions of two leading defense theories, the growth-differentiation balance hypothesis (GDBH) and optimal defense theory (ODT). To achieve this, enemies, fitness components, terpenoids, and protease inhibitors were measured in Solidago altissima and used to construct conventional univariate and structural equation models (SEMs). Leaf-tissue value indices extracted from an SEM revealed a strong correlation between tissue value and terpenoid defense that supports ODT. A tradeoff between serine protease inhibition and growth as well as an indirect tradeoff between growth and terpenoids manifested through galling insects supported the GDBH. Interestingly, there was a strong direct effect of terpenoids on rhizome mass, suggesting service to both storage and defense. The results support established theories but unknown genotypic traits explained much of the variation in defense, confirming the need to integrate emerging theories such as pollination constraints, defense syndromes, tolerance, mutualisms, and facilitation.

Carotenoids are conjugated isoprenoid molecules with many important physiological functions in organisms, including roles in photosynthesis, oxidative stress reduction, vision, diapause, photoperiodism, and immunity. Until recently, it was believed that only plants, microorganisms, and fungi were capable of synthesizing carotenoids and that animals acquired them from their diet, but recent studies have demonstrated that two arthropods (pea aphid and spider mite) possess a pair of genes homologous to those required for the first step of carotenoid biosynthesis. Absent in all other known animal genomes, these genes appear to have been acquired by aphids and spider mites in one or several lateral gene transfer events from a fungal donor. We report the third case of fungal carotenoid biosynthesis gene homologs in an arthropod: flies from the family Cecidomyiidae, commonly known as gall midges. Using phylogenetic analyses we show that it is unlikely that lycopene cyclase/phytoene synthase and phytoene desaturase homologs were transferred singly to an ancient arthropod ancestor; instead we propose that genes were transferred independently from related fungal donors after divergence of the major arthropod lineages. We also examine variation in intron placement and copy number of the carotenoid genes that may underlie function in the midges. This trans-kingdom transfer of carotenoid genes may represent a key innovation, underlying the evolution of phytophagy and plant-galling in gall midges and facilitating their extensive diversification across plant lineages.

The Neotropics harbor an enormous diversity of tachinid flies (Diptera: Tachinidae), yet the fauna remains poorly understood. This is especially true of the tribe Blondeliini, which is particularly diverse in this region and in great need of taxonomic attention. Here, the Neotropical blondeliine genus Erythromelana Townsend is revised. This genus is widely distributed from southern Mexico to northern Argentina, with the Andes being a hotspot of diversity. Known hosts belong to the genus Eois Hübner (Lepidoptera: Geometridae). This revision includes the redescription of three previously described species and the description of 11 new species based on characteristics of adults and immatures. The new species are E. arciforceps sp. nov., E. catarina sp. nov., E. convexiforceps sp. nov., E. cryptica sp. nov., E. curvifrons sp. nov., E. distincta sp. nov., E. ecuadoriana sp. nov., E. eois sp. nov., E. leptoforceps sp. nov., E. napensis sp. nov., and E. woodi sp. nov. A morphological database of 62 characters was constructed to assess morphological variation within and among species and species groups using Principal Components Analysis. Means and medians for these morphological traits were calculated to infer phylogenetic relationships using parsimony. Additionally, a maximum likelihood phylogenetic analysis was performed using COI mtDNA sequences for a subset of eight species. Nominal species E. obscurifrons (Wulp) is treated as a nomen dubium within Erythromelana. Two species previously assigned to Erythromelana appear to represent distinct genera with unclear relationships to this genus and are reinstated as monotypic genera: Myiodoriops marginalis Townsend and Euptilodegeeria obumbrata (Wulp), revived status. Biological and phylogenetic data are used to infer modes of diversification within Erythromelana.

Carotenoids are long conjugated isoprenoid molecules derived mainly from plants and microbial organisms. They are highly diverse, with over 700 identified structures, and are widespread in nature. In addition to their fundamental roles as light-harvesting molecules in photosynthesis, carotenoids serve a variety of functions including visual and colouring pigments, antioxidants and hormone precursors. Although the functions of carotenoids are relatively well studied in plants and vertebrates, studies are severely lacking in insect systems. There is a particular dearth of knowledge on how carotenoids move among trophic levels, influence insect multitrophic interactions and affect evolutionary outcomes. This review explores the known and potential roles that carotenoids and their derivatives have in mediating the ecological interaction of insects with their environment. Throughout the review, we highlight how the fundamental roles of carotenoids in insect physiology might be linked to ecological and evolutionary processes.

Summary 1. Agricultural intensification has led to the removal of semi-wild, perennial vegetation in agricultural landscapes. However, in short-cycle crops, frequent disturbance from insecticides, harvesting and tillage disrupts the establishment of resident communities of natural enemies of pests. Semi-wild perennial vegetation may provide critical habitat for mobile arthropods supporting ecosystem services and sustainable agriculture. 2. We sampled tachinid parasitoids, an important taxon for biological control of vegetable pests, in 35 organic farm fields situated within a mosaic of agricultural, residential and preserved lands in coastal California. Using a GIS, we characterized land-use and vegetative cover within 500 and 1500 m, including grasslands, chaparral, oak woodlands and coniferous forests. 3. The abundance and species richness of tachinid flies captured in Malaise traps in spring and summer were positively associated with the cover of semi-wild perennial vegetation, especially in mesic habitats. The effective number of tachinid species (e H ) was correlated positively with semi-wild perennial vegetation cover and negatively with annual crop cover in the landscape in September and May. 4. The richness of parasitoids emerging from sentinel lepidopteran pests exposed on potted plants within farm fields was negatively associated with annual cropland cover. Parasitism rates dropped precipitously as percentage annual crop cover exceeded species-specific thresholds. 5. Synthesis and applications. Maintaining semi-wild, perennial habitat fragments as refugia to support parasitoids can increase biodiversity and provide ecosystem services in annual and short-cycle crop fields. Our results indicated that crop pests escaped parasitism by two important tachinid species in landscapes with greater than 38% and 51% cover of annual cropland, respectively. Landscape-level research is critical for integrating science and policy to conserve biodiversity, promote sustainable agroecosystems and evaluate new anti-wildlife vegetation removal campaigns that may harm biological control agents while targeting microbial food

1. Adaptive divergence in sympatry is supposed to be inhibited by the homogenizing role of gene flow. However, studies continue to uncover examples of sympatric divergence. In this study, two divergent phenotypes in a complex of four syntopic gall midge morphotypes [nominally Asteromyia carbonifera Osten Saken, Diptera: Cecidomyiidae: Alycaulini] are characterised. The first is a behavioural phenotype governing within-host tissue preference and the second is a trait governing accessory-gland carotenoid quality and quantity. 2. One gall morphotype (crescents) lay most of their eggs on mature tissue whereas the other three gall morphotypes oviposit only on young emerging leaves. Ecological maintenance of this divergent trait appears to be driven by enemy-reduced space. That is, nearly 40% of the crescent morphotype galls that develop high on the plant are attacked by the egg parasitoid Platygaster solidaginis Ashmed, whereas those low on the plant are relatively protected. 3. All morphotypes contain carotenoids in their accessory glands, but the quality and quantity of these pigments differs significantly between the morphotypes and is positively associated with the probability of parasitism by P. solidaginis. 4. Larval salivary glands also contain carotenoids and the plant hormone abscisic acid, which in plants is synthesized from carotenoid precursors and is involved in regulating plant defences. This hormone may facilitate gall development and influence gall morphology. 5. Ecological fitness trade-offs between carotenoids, parasitoid attack, and plant resistance may be fostering adaptive divergence in ovipositional phenotypes and sympatric speciation in this complex of gall midge morphotypes.

This study encompasses three trophic levels, including description and observations of a butterfly caterpillar, its host plant, and two parasitoids, one of which is described here as a new species. We describe the immature stages and larval and adult behaviors of the butterfly Eryphanis greeneyi Penz & DeVries, 2008 (Lepidoptera: Nymphalidae) from the east Andean slopes of Ecuador. We identify Chusquea cf. scandens (Poaceae: Bambusoideae) as the larval food plant and review host associations in the genus Eryphanis and related taxa. We compare the morphology of E. greeneyi caterpillars to related taxa feeding on Chusquea and discuss the function of anal combs and associated frass throwing behavior. We describe a new species of braconid wasp Protapanteles eryphanidis sp. n. Whitfield and a tachinid fly, Winthemia sp. nr. analis, reared from field-collected larvae of E. greeneyi and evaluate their taxonomic position and association with E. greeneyi.

We analysed the canopy and understorey communities of flies in the family Tachinidae, the most diverse group of parasitoid Diptera, in a small and iso- lated temperate plain forest in northern Italy. Our objective was to assess whether and how these communities differ from one another, and how species distribution relates to forest structure, host distribution, mating sites, and season. 2. The study was carried out in 2008 with 14 Malaise traps installed between April and November in an equal number of sites randomly selected inside the forest, seven on the ground and seven in the tree canopy. 3. Overall species richness, abundance, and turnover were greater in the understo- rey traps, but most diversity metrics indicate greater overall diversity and evenness in the canopy traps. Community ordination and estimates of beta diversity indicate that the two habitat-associated communities are distinct and should both be consid- ered in assessments of insect diversity and community structure. Indicator species values revealed the presence of a number of species that were effective indicators of canopy and understorey habitats. No strong male bias in canopy traps was observed across species; however, the only significant sex ratio biases in the canopy were towards males. Both male and female biases were observed in understorey traps, depending upon the species.

Although tachinids are one of the most diverse families of Diptera and represent the largest group of nonhymenopteran parasitoids, their local diversity and distribution patterns of most species in the family are poorly known. In this study, 2 yr of samples from a Malaise trap were used to analyze the local richness and temporal distribution of a tachinid community in southwestern Ohio. In total, 883 tachinid specimens were collected, consisting of 117 species belonging to 69 genera. The majority of the specimens were collected in fall, followed by summer and spring, with 67, 22, and 11%, respectively. Estimated rarefaction curves indicate a predicted species richness of 217 species and suggest that we have sampled only a fraction of the diversity of Tachinidae occurring at this particular site. The species recorded in this study represent 16 and 19% of the species that are likely to occur in the northeastern United States and Ohio, respectively. In North America, there have been few, if any, previous attempts to quantify the diversity of tachinids on a local scale. Our results provide a baseline for understanding the temporal and spatial diversity of these ecologically and agronomically important parasitoids.

The Ambrosia gall midge [Asteromyia carbonifera (Osten Sacken) (Diptera: Cecidomyiidae: Alycaulini)] consists, in part, of a complex of genetically differentiated populations that have diverged in gall morphology on the host plant Solidago altissima L. (Asteraceae). This divergence appears to be an incipient adaptive radiation that may be driven by parasitoid pressure. Understanding the mechanisms driving this genetic and phenotypic diversification requires a close examination of the relationship between the midge and its fungal associate Botryosphaeria dothidea (Moug.) Ces. & De Not. (Ascomycota: Dothideomycetes), whose mycelia actually form the protective gall structure. We used manipulative experiments to test the degree of interdependency of the fungus and the midge, and we employed field and laboratory studies to gain insight into the source of fungal conidia, which our data and observations indicate are collected by females and stored in specialized pockets (mycangia) on the ovipositor. Manipulative experiments demonstrate that fungal proliferation on the host plant is dependent on the midge larvae and larvae exhibit significant growth on the fungus alone. Field observations and experiments were unable to identify the source of mycangial conidia; however, analyses of conidia shape suggest a biotrophic source. We conclude that this association is an obligatory mutualism with respect to successful gall formation. These findings corroborate recent findings that the primary food source of the midge is the gall fungus.

Multitrophic interactions play key roles in the origin and maintenance of species diversity, and the study of these interactions has contributed to important theoretical advances in ecology and evolutionary biology. Nevertheless, most biodiversity inventories focus on static species lists, and prominent theories of diversity still ignore trophic interactions. The lack of a simple interaction metric that is analogous to species richness is one reason why diversity of interactions is not examined as a response or predictor variable in diversity studies. Using plant–herbivore–enemy trophic chains as an example, we develop a simple metric of diversity in which richness, diversity indices (e.g., Simpson's 1/D), and rarefaction diversity are calculated with links as the basic unit rather than species. Interactions include all two-link (herbivore–plant and enemy–herbivore) and three-link (enemy–herbivore–plant) chains found in a study unit. This metric is different from other indices, such as traditional diversity measures, connectivity and interaction diversity in food-web studies, and the diversity of interaction index in behavioral studies, and it is easier to compute. Using this approach to studying diversity provides novel insight into debates about neutrality and correlations between diversity, stability, productivity, and ecosystem services. RESUMEN Las interacciones multitroficas son en gran parte responsables por el origen y el mantenimiento de la diversidad de especies, y las investigaciones de estas interacciones han contribuido a avances teoricos en ecologia y biologia evolutiva. Sin embargo, en la mayoria de los inventarios de biodiversidad se hacen listas estaticas de especies, y las teorias de la diversidad mas prominentes siguen ignorando las interacciones troficas. La falta de una metrica de interaccion sencilla y que sea analoga a la riqueza de especies es una de las razones para que no se examine la diversidad de interacciones como una respuesta o una variable dependiente en estudios de diversidad. Usamos las cadenas troficas de plantas-herbivoros-enemigos como un ejemplo y desarrollamos una variable simple de diversidad en la cual se puede calcular la riqueza de interacciones o los indices de diversidad (por ejemplo, 1 / D de Simpson) y la diversidad de rarefaccion; se usan los enlaces como la unidad basica en lugar de las especies. Las interacciones incluyen todas las interacciones de dos enlaces (planta-herbivoro y herbivoro-enemigo) y de tres enlaces (enemigo-herbivoro-planta) que se encuentra en muestreos o censos por transectos o por parcelas. Esta variable es diferente de otros indices de diversidad tradicionalmente usados como la conectividad de interaccion en estudios de cadenas troficas, y el indice de diversidad de interacciones que se usa en estudios comportamentales, ademas es mas facil de calcular. Usando esta aproximacion para estudiar la diversidad de interacciones creara nuevas perspectivas en debates sobre la teoria neutral y las correlaciones entre la diversidad, la estabilidad, la productividad, y servicios de los ecosistemas.

Gall-forming insects provide ideal systems to analyze the evolution of host-parasite interactions and understand the ecological interactions that contribute to evolutionary diversification. Flies in the family Cecidomyiidae represent the largest radiation of gall-forming insects and are characterized by complex trophic interactions with plants, fungal symbionts, and predators. We analyzed the phylogenetic history and evolutionary associations of the North American cecidomyiid genus Asteromyia, which is engaged in a complex and perhaps co-evolving community of interactions with host-plants, fungi, and parasitoids. Mitochondrial gene trees generally support current classifications, but reveal extensive cryptic diversity within the eight named species. Asteromyia likely radiated after their associated host-plants in the Astereae, but species groups exhibit strong associations with specific lineages of Astereae. Evolutionary associations with fungal mutualists are dynamic, however, and suggest rapid and perhaps coordinated changes across trophic levels.

Here we describe the immature stages and ecological associations of Antirrhea adoptiva porphyrosticta Watkins, 1928 (Lepidoptera:Nymphalidae:Morphinae). The cloud forest bamboo, Chusquea scandens Kunth (Bambusoidea: Poaceae), serves as the larval food plant for this butterfly in eastern Ecuador, the first hostplant record for Antirrhea outside the family Arecaceae. The larvae of A. adoptiva porphyrosticta are superficially similar to those of other Antirrhea species. We also provide observations on adult and larval behavior. Caterpillars of this butterfly species are parasitized by tachinid flies, as well as by Ichneumonidae and a newly described braconid wasp.

Most of the unknown biological diversity of macro-organisms remaining to be discovered and described lies in the tropical regions of the world and consists primarily of insects. Those insects with parasitoid lifestyles constitute a significant portion of insect diversity, yet parasitoids are among the most poorly known of major insect guilds in the humid tropics. Here we describe and analyze the richness of one diverse taxon of parasitoids, flies in the family Tachinidae, reared from Lepidoptera as part of a biological survey of Lepidoptera and their parasitoids in one mid-elevation (2000 m) area in the northeast Ecuadorian Andes. One hundred fifty-seven separable tachinid "morpho-species" were reared from approximately 160 species of Lepidoptera in 16 families. These tachinid flies were recovered from a sample of over 12,800 successful caterpillar rearing events that resulted in either adult Lepidoptera or parasitoids. Tachinid species accumulation and rarefaction curves exhibit no sign of reaching an asymptote and richness estimators indicate that the community likely consists of nearly twice this number of species (at minimum). Most tachinid species were reared infrequently, with 50% being represented by a single individual. The majority of species appeared to be relatively specialized on one or a few related hosts, but sampling was insufficient to make strong inferences regarding host range. The tribes Blondeliini and Goniini were the best represented, but some tribes that were expected to be common such as Tachinini and Winthemiini were poorly represented. The estimates of tachinid species richness derived here are suggestive of a far more diverse tachinid community than in temperate localities in North America. Additional rearing of Lepidoptera, as well as other herbivorous insect taxa, along with the use of additional collecting methods will be necessary to achieve a more accurate understanding of the richness of tropical Tachinidae and their contribution to broader patterns of tropical biodiversity.

Many species of Tachinidae are important parasitoids of herbivorous insects in natural and managed systems; yet, little is known about tachinid diversity and how this diversity is distributed across space and time. Here, pan trap sampling was used to analyze the richness, microhabitat speciÞcity, and seasonal diversity of a tachinid parasitoid community in an oakÐ mesquite savanna of Southeastern Arizona. Twenty-four traps were set out monthly during the growing season in three different microhabitats (open grassland, woodland understory, and woodland canopy). In total, 79 tachinid species were sampled with an estimated total diversity of 122 species. Most individual traps sampled few species; yet, variation in species composition ( diversity) among sampling dates and microhabitats was high, accounting for 40 Ð70% of the total diversity. SigniÞcant intraspeciÞc aggregation was not observed across traps or microhabitats, but it was observed across dates, suggesting that the activity of tachinid species may be associated with host phenology and seasonal periods of precipitation. The diversity associated with microhabitat and sampling date was signiÞcantly greater than expected. Tachinid species diversity was highest in the canopy traps, whereas the open-exposed traps exhibited high abundances of relatively few species, and understory traps sampled few individuals of few species. Most species tended to be sampled where their hosts would be expected to be found, although males were also frequently sampled in microhabitats associated with mate Þnding. These patterns of diversity and abundance may aid in understanding parasitoidÐ host associations and variation in rates of parasitism by tachinid sies.

There is growing realization that intraspecific polyploidy in plants has important implications for the evolution of plant diversity and for plant and plant-herbivore community dynamics. Nonetheless, the rate at which polyploid cytotypes arise and their fate in local mixed-cytotype populations are not well understood. Although within- and especially among-population cytotype variation has been documented in many plants, particularly at the largest (continental) spatial scales, local and regional distributions of cy totypes have been well characterized only for a handful of species. Furthermore, few if any studies have addressed both local and regional patterns in the same plant species with sufficient power to detect cytotype variation on both scales. We assessed the co occurrence and frequency distributions of diploid, tetraploid, and hexaploid cytotypes of Solidago altissima (Asteraceae) at local and regional spatial scales, sampling across a zone of presumed ploidy-form overlap in the midwestern United States. Determining ploidy by flow cytometry, we found many local populations containing two or three cytotypes, with the relative frequencies of cytotypes varying dramatically even among neighboring populations. AFLP marker data suggest that polyploid cytotypes of S. altissima likely have multiple origins from different diploid lineages. Our results are consistent with recent perspectives on polyploidization as an evolutionarily dynamic phenomenon with frequent multiple origins of higher-ploidy cytotypes.