Using DNA-barcoded Malaise trap samples to measure impact of a geothermal energy project on the biodiversity of a Costa Rican old-growth rain forest.

We report one year (2013-2014) of biomonitoring an insect community in a tropical old-growth rain forest, during construction of an industrial-level geothermal electricity project. This is the first-year reaction by the species-rich insect biodiversity; six subsequent years are being analyzed now. The site is on the margin of a UNESCO Natural World Heritage Site, Área de Conservación Guanacaste (ACG), in northwestern Costa Rica. This biomonitoring is part of Costa Rica's ongoing efforts to sustainably retain its wild biodiversity through biodevelopmental integration with its societies. Essential tools are geothermal engineering needs, entomological knowledge, insect species-rich forest, government-NGO integration, common sense, DNA barcoding for species-level identification, and Malaise traps. This research is tailored for integration with its society at the product level. We combine an academic view with on-site engineering decisions. This biomonitoring requires alpha-level DNA barcoding combined with centuries of morphology-based entomological taxonomy and ecology. Not all desired insect community analyses are performed; they are for data from subsequent years combined with this year. We provide enough analysis to be used by both guilds now. This biomonitoring has shown, for the first year, that the geothermal project impacts only the biodiversity within a zone less than 50 m from the project margin.

Bromeliads affect the interactions and composition of invertebrates on their support tree.

Individual species can have profound effects on ecological communities, but, in hyperdiverse systems, it can be challenging to determine the underlying ecological mechanisms. Simplifying species' responses by trophic level or functional group may be useful, but characterizing the trait structure of communities may be better related to niche processes. A largely overlooked trait in such community-level analyses is behaviour. In the Neotropics, epiphytic tank bromeliads (Bromeliaceae) harbour a distinct fauna of terrestrial invertebrates that is mainly composed of predators, such as ants and spiders. As these bromeliad-associated predators tend to forage on the bromeliads' support tree, they may influence the arboreal invertebrate fauna. We examined how, by increasing associated predator habitat, bromeliads may affect arboreal invertebrates. Specifically, we observed the trophic and functional group composition, and the behaviour and interspecific interactions of arboreal invertebrates in trees with and without bromeliads. Bromeliads modified the functional composition of arboreal invertebrates, but not the overall abundance of predators and herbivores. Bromeliads did not alter the overall behavioural profile of arboreal invertebrates, but did lead to more positive interactions in the day than at night, with a reverse pattern on trees without bromeliads. In particular, tending behaviours were influenced by bromeliad-associated predators. These results indicate that detailed examination of the functional affiliations and behaviour of organisms can reveal complex effects of habitat-forming species like bromeliads, even when total densities of trophic groups are insensitive.

Species-level predation network uncovers high prey specificity in a Neotropical army ant community.

Army ants are among the top arthropod predators and considered keystone species in tropical ecosystems. During daily mass raids with many thousand workers, army ants hunt live prey, likely exerting strong top-down control on prey species. Many tropical sites exhibit a high army ant species diversity (>20 species), suggesting that sympatric species partition the available prey niches. However, whether and to what extent this is achieved has not been intensively studied yet. We therefore conducted a large-scale diet survey of a community of surface-raiding army ants at La Selva Biological Station in Costa Rica. We systematically collected 3,262 prey items from eleven army ant species (genera Eciton, Nomamyrmex and Neivamyrmex). Prey items were classified as ant prey or non-ant prey. The prey nearly exclusively consisted of other ants (98%), and most booty was ant brood (87%). Using morphological characters and DNA barcoding, we identified a total of 1,103 ant prey specimens to the species level. One hundred twenty-nine ant species were detected among the army ant prey, representing about 30% of the known local ant diversity. Using weighted bipartite network analyses, we show that prey specialization in army ants is unexpectedly high and prey niche overlap very small. Besides food niche differentiation, we uncovered a spatiotemporal niche differentiation in army ant raid activity. We discuss competition-driven multidimensional niche differentiation and predator-prey arms races as possible mechanisms underlying prey specialization in army ants. By combining systematic prey sampling with species-level prey identification and network analyses, our integrative approach can guide future research by portraying how predator-prey interactions in complex communities can be reliably studied, even in cases where morphological prey identification is infeasible.

DNA barcoding reveals diversity of Hymenoptera and the dominance of parasitoids in a sub-arctic environment.

BACKGROUND: Insect diversity typically declines with increasing latitude, but previous studies have shown conflicting latitude-richness gradients for some hymenopteran parasitoids. However, historical estimates of insect diversity and species richness can be difficult to confirm or compare, because they may be based upon dissimilar methods. As a proxy for species identification, we used DNA barcoding to identify molecular operational taxonomic units (MOTUs) for 7870 Hymenoptera specimens collected near Churchill, Manitoba, from 2004 through 2010. RESULTS: We resolved 1630 MOTUs for this collection, of which 75% (1228) were ichneumonoids (Ichneumonidae + Braconidae) and 91% (1484) were parasitoids. We estimate the total number of Hymenoptera MOTUs in this region at 2624-2840. CONCLUSIONS: The diversity of parasitoids in this sub-Arctic environment implies a high diversity of potential host species throughout the same range. We discuss these results in the contexts of resolving interspecific interactions that may include cryptic species, and developing reproducible methods to estimate and compare species richness across sites and between surveys, especially when morphological specialists are not available to identify every specimen.

Revision of Belvosia Robineau-Desvoidy (Diptera, Tachinidae) and 33 new species from Area de Conservación Guanacaste in northwestern Costa Rica with a key to known North and Mesoamerican species.

Background: This revision is part of a continuing series of taxonomic work aimed at the description of new taxa and the redescription of known taxa of the Tachinidae of Area de Conservación Guanacaste in northwestern Costa Rica. Here we describe 33 new species in the genus Belvosia Robineau-Desvoidy, 1830 (Diptera: Tachinidae). All species described here were reared from this ongoing inventory of wild-caught caterpillars spanning a variety of families (Lepidoptera: Erebidae, Eupterotidae, Noctuidae, Notodontidae, Saturniidae, and Sphingidae). We provide a morphological description of each species with limited information on life history, molecular data, and photographic documentation. In addition to the new species, the authors provide a redescription of the genus Belvosia, as well as provide a key to the identification of the species present in the Meso- and North-American fauna. New information: The following 33 new species of Belvosia Robineau-Desvoidy, 1830, all authored by Fleming & Woodley, are described: Belvosiaadrianguadamuzi Fleming & Woodley sp. n., Belvosiaanacarballoae Fleming & Woodley sp. n., Belvosiaangelhernandezi Fleming & Woodley sp. n., Belvosiabrigittevilchezae Fleming & Woodley sp. n., Belvosiaalixtomoragai Fleming & Woodley sp. n., Belvosiacarolinacanoae Fleming & Woodley sp. n., Belvosiaciriloumanai Fleming & Woodley sp. n., Belvosiadiniamartinezae Fleming & Woodley sp. n., Belvosiaduniagarciae Fleming & Woodley sp. n., Belvosiaduvalierbricenoi Fleming & Woodley sp. n., Belvosiaeldaarayae Fleming & Woodley sp. n., Belvosiaeliethcantillanoae Fleming & Woodley sp. n., Belvosiafreddyquesadai Fleming & Woodley sp. n., Belvosiagloriasihezarae Fleming & Woodley sp. n., Belvosiaguillermopereirai Fleming & Woodley sp. n., Belvosiaharryramirezi Fleming & Woodley sp. n., Belvosiahazelcambroneroae Fleming & Woodley sp. n., Belvosiajorgehernandezi Fleming & Woodley sp. n., Belvosiajosecortezi Fleming & Woodley sp. n., Belvosiajoseperezi Fleming & Woodley sp. n., Belvosiakeinoraragoni Fleming & Woodley sp. n., Belvosialuciariosae Fleming & Woodley sp. n., Belvosiamanuelpereirai Fleming & Woodley sp. n., Belvosiamanuelriosi Fleming & Woodley sp. n., Belvosiaminorcarmonai Fleming & Woodley sp. n., Belvosiaosvaldoespinozai Fleming & Woodley sp. n., Belvosiapabloumanai Fleming & Woodley sp. n., Belvosiapetronariosae Fleming & Woodley sp. n., Belvosiaricardocaleroi Fleming & Woodley sp. n., Belvosiarobertoespinozai Fleming & Woodley sp. n., Belvosiarostermoragai Fleming & Woodley sp. n., Belvosiaruthfrancoae Fleming & Woodley sp. n., Belvosiasergioriosi Fleming & Woodley sp. n.Belvosiacanalis Aldrich, 1928 is reared and recorded from the inventory; new information relative to host is provided and the species is rediscribed.The following are proposed by Fleming & Woodley as new synonyms of Belvosia Robineau-Desvoidy, 1830: Brachybelvosia Townsend, 1927 syn. n., Belvosiomimops Townsend, 1935 syn. n.The following three new combinations are proposed as a result of the new synonymies: Belvosiabrasilensis (Townsend, 1927), comb. n.; and Belvosiabarbiellinii (Townsend, 1935), comb. n.The authors also propose the following new synonymies: Belvosiabrasilensis (Townsend, 1927) = Belvosiaaurulenta (Bigot, 1888), syn. n.; Belvosiapollinosa Rowe, 1933 = Belvosiaborealis Aldrich, 1928 syn. n.; Belvosiaweyenberghiana (Wulp, 1883) = Belvosiafuliginosa (Walker, 1853) syn. n.; Belvosiabrasiliensis Townsend, 1927 = Belvosiafuliginosa (Walker, 1853) syn. n.; Belvosialuteola Coquillett, 1900 = Belvosiaochriventris (Wulp, 1890) syn. n.; Belvosiasocia (Walker, 1853) = Belvosiaproxima (Walker, 1853) syn. n.; Belvosiachrysopyga (Bigot, 1887) = Belvosiaunifasciata (Robineau-Desvoidy, 1830) syn. n.; Belvosiachrysopygata (Bigot, 1888) = Belvosiaunifasciata (Robineau-Desvoidy, 1830) syn. n.

Amphibian diversity across three adjacent ecosystems in Área de Conservación Guanacaste, Costa Rica.

Amphibians are the most threatened species-rich vertebrate group, with species extinctions and population declines occurring globally, even in protected and seemingly pristine habitats. These 'enigmatic declines' are generated by climate change and infectious diseases. However, the consequences of these declines are undocumented as no baseline ecological data exists for most affected areas. Like other neotropical countries, Costa Rica, including Área de Conservación Guanacaste (ACG) in north-western Costa Rica, experienced rapid amphibian population declines and apparent extinctions during the past three decades. To delineate amphibian diversity patterns within ACG, a large-scale comparison of multiple sites and habitats was conducted. Distance and time constrained visual encounter surveys characterised species richness at five sites-Murciélago (dry forest), Santa Rosa (dry forest), Maritza (mid-elevation dry-rain forest intersect), San Gerardo (rainforest) and Cacao (cloud forest). Furthermore, species-richness patterns for Cacao were compared with historic data from 1987-8, before amphibians declined in the area. Rainforests had the highest species richness, with triple the species of their dry forest counterparts. A decline of 45% (20 to 11 species) in amphibian species richness was encountered when comparing historic and contemporary data for Cacao. Conservation efforts sometimes focus on increasing the resilience of protected areas, by increasing their range of ecosystems. In this sense ACG is unique containing many tropical ecosystems compressed in a small geographic space, all protected and recognised as a UNESCO world heritage site. It thus provides an extraordinary platform to understand changes, past and present, and the resilience of tropical ecosystems and assemblages, or lack thereof, to climate change.

A revision of the parasitoid wasp genus Alphomelon Mason with the description of 30 new species (Hymenoptera, Braconidae).

The parasitoid wasp genus Alphomelon Mason, 1981 is revised, based on a combination of basic morphology (dichotomous key and brief diagnostic descriptions), DNA barcoding, biology (host data and wasp cocoons), and distribution data. A total of 49 species is considered; the genus is almost entirely Neotropical (48 species recorded from that region), but three species reach the Nearctic, with one of them extending as far north as 45° N in Canada. Alphomelon parasitizes exclusively Hesperiinae caterpillars (Lepidoptera: Hesperiidae), mostly feeding on monocots in the families Arecaceae, Bromeliaceae, Cannaceae, Commelinaceae, Heliconiaceae, and Poaceae. Most wasp species parasitize either on one or very few (2-4) host species, usually within one or two hesperiine genera; but some species can parasitize several hosts from up to nine different hesperiine genera. Among species with available data for their cocoons, roughly half weave solitary cocoons (16) and half are gregarious (17); cocoons tend to be surrounded by a rather distinctive, coarse silk (especially in solitary species, but also distinguishable in some gregarious species). Neither morphology nor DNA barcoding alone was sufficient on its own to delimit all species properly; by integrating all available evidence (even if incomplete, as available data for every species is different) a foundation is provided for future studies incorporating more specimens, especially from South America. The following 30 new species are described: cruzi, itatiaiensis, and palomae, authored by Shimbori & Fernandez-Triana; and adrianguadamuzi, amazonas, andydeansi, calixtomoragai, carolinacanoae, christerhanssoni, diniamartinezae, duvalierbricenoi, eldaarayae, eliethcantillanoae, gloriasihezarae, guillermopereirai, hazelcambroneroae, josecortesi, keineraragoni, luciarosae, manuelriosi, mikesharkeyi, osvaldoespinozai, paramelanoscelis, paranigriceps, petronariosae, ricardocaleroi, rigoi, rostermoragai, sergioriosi, and yanayacu, authored by Fernandez-Triana & Shimbori.

Do bromeliads affect the arboreal ant communities on orange trees in northwestern Costa Rica?

Ants often interact with other invertebrates as predators or mutualists. Epiphytic bromeliads provide nesting sites for ants, and could increase ant abundances in the tree canopy. We surveyed ants in the foliage of orange trees that either hosted bromeliads or did not. To determine if observed associations between bromeliads and tree ants were causal, we removed bromeliads from half of the trees, and resurveyed ants six weeks later. Our results show that bromeliad presence is correlated with higher ant abundances and different species of ants on orange trees during the dry season. This increase in ant abundance was driven primarily by Solenopsis ants, which were both numerous and found to facultatively nest in bromeliads. Bromeliad removal did not affect either ant abundance or composition, potentially because this manipulation coincided with the transition from dry to wet season. Other ant species were never encountered nesting in bromeliads, and the abundances of such ants on tree leaves were unaffected by bromeliad presence or removal. Considering the importance of ants in herbivore regulation, our findings suggest that bromeliads-through their association with ants-could indirectly be associated with biological control in agricultural systems.

Frequency, topic, and preferences: Tracking student engagement with several modalities of student-instructor contact in a first-year course.

Meaningful student-instructor interactions during an undergraduate degree course can have important effects on student learning. The format by which those interactions are made possible can vary greatly. We investigated the preferred modality of contact and students' reasons for contact across several modalities in a first-year biology course. We tracked student-instructor contact for two-course instructors who team teach collaboratively (rather than sequentially) across two-course sections. Both instructors had identical scores on student evaluations of approachability. Student-instructor contact was facilitated using five 'student hour' modalities: (a) in office by appointment, (b) 1 h per week, in office drop in, (c) 1 h per week, virtual chat, (d) by email, (e) 10 min immediately after class. Though email was the preferred method of contact, the period immediately following the class instruction was the most popular of the face-to-face options. We note significant differences in the distribution of workload across the two instructors and make recommendations for increasing the accessibility of student-instructor contact and for equity in workload to support student learning.

More discussion of minimalist species descriptions and clarifying some misconceptions contained in Meier et al. 2021.

This is a response to a preprint version of "A re-analysis of the data in Sharkey et al.'s (2021) minimalist revision reveals that BINs do not deserve names, but BOLD Systems needs a stronger commitment to open science", https://www.biorxiv.org/content/10.1101/2021.04.28.441626v2. Meier et al. strongly criticized Sharkey et al.'s publication in which 403 new species were deliberately minimally described, based primarily on COI barcode sequence data. Here we respond to these criticisms. The following points are made: 1) Sharkey et al. did not equate BINs with species, as demonstrated in several examples in which multiple species were found to be in single BINs. 2) We reiterate that BINs were used as a preliminary sorting tool, just as preliminary morphological identification commonly sorts specimens based on color and size into unit trays; despite BINs and species concepts matching well over 90% of species, this matching does not equate to equality. 3) Consensus barcodes were used only to provide a diagnosis to conform to the rules of the International Code of Zoological Nomenclature just as consensus morphological diagnoses are. The barcode of a holotype is definitive and simply part of its cellular morphology. 4) Minimalist revisions will facilitate and accelerate future taxonomic research, not hinder it. 5) We refute the claim that the BOLD sequences of Plesiocoelusvanachterbergi are pseudogenes and demonstrate that they simply represent a frameshift mutation. 6) We reassert our observation that morphological evidence alone is insufficient to recognize species within species-rich higher taxa and that its usefulness lies in character states that are congruent with molecular data. 7) We show that in the cases in which COI barcodes code for the same amino acids in different putative species, data from morphology, host specificity, and other ecological traits reaffirm their utility as indicators of genetically distinct lineages.

Cryptic biodiversity of tropical hesperiid caterpillar-attacking parasitoid wasps: three new species of Creagrura Townes (Hymenoptera, Ichneumonidae, Cremastinae) from Costa Rica and Perú.

Background: We describe three new species of the previously monotypic genus Creagrura Townes from Central and South America: C.alejandromasisi sp. n. and C.rogerblancoi sp. n. from Costa Rica and C.allpahuaya sp. n. from Peru, all of which emphasise the unknown parasitoid insect diversity yet to be revealed in the tropics. New information: Host relationships of the two Costa Rican species are described in detail. In addition, it is inferred that the Creagrura wasps find and oviposit in the caterpillar when it is exposed at night, rather than when it is concealed during daylight hours.

Comparison of traditional and DNA metabarcoding samples for monitoring tropical soil arthropods (Formicidae, Collembola and Isoptera).

The soil fauna of the tropics remains one of the least known components of the biosphere. Long-term monitoring of this fauna is hampered by the lack of taxonomic expertise and funding. These obstacles may potentially be lifted with DNA metabarcoding. To validate this approach, we studied the ants, springtails and termites of 100 paired soil samples from Barro Colorado Island, Panama. The fauna was extracted with Berlese-Tullgren funnels and then either sorted with traditional taxonomy and known, individual DNA barcodes ("traditional samples") or processed with metabarcoding ("metabarcoding samples"). We detected 49 ant, 37 springtail and 34 termite species with 3.46 million reads of the COI gene, at a mean sequence length of 233 bp. Traditional identification yielded 80, 111 and 15 species of ants, springtails and termites, respectively; 98%, 37% and 100% of these species had a Barcode Index Number (BIN) allowing for direct comparison with metabarcoding. Ants were best surveyed through traditional methods, termites were better detected by metabarcoding, and springtails were equally well detected by both techniques. Species richness was underestimated, and faunal composition was different in metabarcoding samples, mostly because 37% of ant species were not detected. The prevalence of species in metabarcoding samples increased with their abundance in traditional samples, and seasonal shifts in species prevalence and faunal composition were similar between traditional and metabarcoding samples. Probable false positive and negative species records were reasonably low (13-18% of common species). We conclude that metabarcoding of samples extracted with Berlese-Tullgren funnels appear suitable for the long-term monitoring of termites and springtails in tropical rainforests. For ants, metabarcoding schemes should be complemented by additional samples of alates from Malaise or light traps.

Molecular analysis of parasitoid linkages (MAPL): gut contents of adult parasitoid wasps reveal larval host.

Metamorphosing insects often have complex and poorly known life histories. In particular, what they feed on during their larval stages remains unknown for the vast majority of species, and its documentation only results from difficult and time-intensive field observations, rearing or dissections. Through the application of a DNA analysis of gut contents in adult parasitoid wasps, we were able to selectively sequence a diagnostic DNA marker that permitted the identification of the host used by these wasps during their larval stages. By reproducing these results in species with different life histories, we excluded other potential sources of host DNA, confirming that after ingestion by the parasitoid larva the host DNA can persist through metamorphosis in the abdominal contents of the adult wasp. Our discovery considerably extends the applicability of molecular analysis of gut contents by enabling the documentation of food used by insects during their larval stages and thus increasing the accuracy and precision of food web studies. The 24% success rate of our approach is surprisingly high considering the challenging context for host DNA preservation, and we discuss the factors possibly affecting this rate. We propose molecular analysis of parasitoid linkages (MAPL) as a new method to document host-parasitoid associations at a faster pace and with unrivalled precision. Because of the key regulatory role of parasitoid wasps in ecosystems, which makes them the most commonly used biological control agents, MAPL will have immediate applications in both basic and applied biological sciences.

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.

Revison of Metaplagia Coquillett (Diptera: Tachinidae) with description of five new species from Area de Conservación Guanacaste in northwestern Costa Rica.

BACKGROUND: We revise the genus Metaplagia Coquillett, 1895 and describe five new species from Area de Conservación Guanacaste (ACG) in northwestern Costa Rica. All new species were reared from an ongoing inventory of wild-caught caterpillars spanning a variety of species within the family Sphingidae (Lepidoptera: Sphingidae). Our study provides a concise description of each new species using morphology, life history, molecular data and photographic documentation. In addition to the new species, the authors provide a re-description of the genus and a revised key to the species of Metaplagia. NEW INFORMATION: The following five new species of Metaplagia are described: Metaplagia leahdennisae Fleming & Wood sp. n., Metaplagia lindarobinsonae Fleming & Wood sp. n., Metaplagia paulinesaribasae Fleming & Wood sp. n., Metaplagia robinsherwoodae Fleming & Wood sp. n. and Metaplagia svetlanakozikae Fleming & Wood sp. n.The following is proposed by Fleming & Wood as new combination of Plagiomima Brauer & Bergenstamm, 1891: Plagiomima latifrons (Reinhard, 1956) comb. n.

Torix Rickettsia are widespread in arthropods and reflect a neglected symbiosis.

BACKGROUND: Rickettsia are intracellular bacteria best known as the causative agents of human and animal diseases. Although these medically important Rickettsia are often transmitted via haematophagous arthropods, other Rickettsia, such as those in the Torix group, appear to reside exclusively in invertebrates and protists with no secondary vertebrate host. Importantly, little is known about the diversity or host range of Torix group Rickettsia. RESULTS: This study describes the serendipitous discovery of Rickettsia amplicons in the Barcode of Life Data System (BOLD), a sequence database specifically designed for the curation of mitochondrial DNA barcodes. Of 184,585 barcode sequences analysed, Rickettsia is observed in ∼0.41% of barcode submissions and is more likely to be found than Wolbachia (0.17%). The Torix group of Rickettsia are shown to account for 95% of all unintended amplifications from the genus. A further targeted PCR screen of 1,612 individuals from 169 terrestrial and aquatic invertebrate species identified mostly Torix strains and supports the "aquatic hot spot" hypothesis for Torix infection. Furthermore, the analysis of 1,341 SRA deposits indicates that Torix infections represent a significant proportion of all Rickettsia symbioses found in arthropod genome projects. CONCLUSIONS: This study supports a previous hypothesis that suggests that Torix Rickettsia are overrepresented in aquatic insects. In addition, multiple methods reveal further putative hot spots of Torix Rickettsia infection, including in phloem-feeding bugs, parasitoid wasps, spiders, and vectors of disease. The unknown host effects and transmission strategies of these endosymbionts make these newly discovered associations important to inform future directions of investigation involving the understudied Torix Rickettsia.

Minimalist revision and description of 403 new species in 11 subfamilies of Costa Rican braconid parasitoid wasps, including host records for 219 species.

Three new genera are described: Michener (Proteropinae), Bioalfa (Rogadinae), and Hermosomastax (Rogadinae). Keys are given for the New World genera of the following braconid subfamilies: Agathidinae, Braconinae, Cheloninae, Homolobinae, Hormiinae, Ichneutinae, Macrocentrinae, Orgilinae, Proteropinae, Rhysipolinae, and Rogadinae. In these subfamilies 416 species are described or redescribed. Most of the species have been reared and all but 13 are new to science. A consensus sequence of the COI barcodes possessed by each species is employed to diagnose the species, and this approach is justified in the introduction. Most descriptions consist of a lateral or dorsal image of the holotype, a diagnostic COI consensus barcode, the Barcode Index Number (BIN) code with a link to the Barcode of Life Database (BOLD), and the holotype specimen information required by the International Code of Zoological Nomenclature. The following species are treated and those lacking authorship are newly described here with authorship attributable to Sharkey except for the new species of Macrocentrinae which are by Sharkey & van Achterberg: AGATHIDINAE: Aerophiluspaulmarshi, Mesocoelusdavidsmithi, Neothlipsisbobkulai, Plesiocoelusvanachterbergi, Pneumagathiserythrogastra (Cameron, 1905), Therophilusbobwhartoni, T.donaldquickei, T.gracewoodae, T.maetoi, T.montywoodi, T.penteadodiasae, Zacremnopsbrianbrowni, Z.coatlicue Sharkey, 1990, Zacremnopscressoni (Cameron, 1887), Z.ekchuah Sharkey, 1990, Z.josefernandezi, Zelomorphasarahmeierottoae. BRACONINAE: Braconalejandromarini, B.alejandromasisi, B.alexamasisae, B.andresmarini, B.andrewwalshi, B.anniapicadoae, B.anniemoriceae, B.barryhammeli, B.bernardoespinozai, B.carlossanabriai, B.chanchini, B.christophervallei, B.erasmocoronadoi, B.eugeniephillipsae, B.federicomatarritai, B.frankjoycei, B.gerardovegai, B.germanvegai, B.isidrochaconi, B.jimlewisi, B.josejaramilloi, B.juanjoseoviedoi, B.juliodiazi, B.luzmariaromeroae, B.manuelzumbadoi, B.marialuisariasae, B.mariamartachavarriae, B.mariorivasi, B.melissaespinozae, B.nelsonzamorai, B.nicklaphami, B.ninamasisae, B.oliverwalshi, B.paulamarinae, B.rafamoralesi, B.robertofernandezi, B.rogerblancoi, B.ronaldzunigai, B.sigifredomarini, B.tihisiaboshartae, B.wilberthbrizuelai, Digonogastramontylloydi, D.montywoodi, D.motohasegawai, D.natwheelwrighti, D.nickgrishini. CHELONINAE: Adeliusadrianguadamuzi, A.gauldi Shimbori & Shaw, 2019, A.janzeni Shimbori & Shaw, 2019, Ascogastergloriasihezarae, A.grettelvegae, A.guillermopereirai, A.gustavoecheverrii, A.katyvandusenae, A.luisdiegogomezi, Chelonusalejandrozaldivari, C.gustavogutierrezi, C.gustavoinduni, C.harryramirezi, C.hartmanguidoi, C.hazelcambroneroae, C.iangauldi, C.isidrochaconi, C.janecheverriae, C.jeffmilleri, C.jennyphillipsae, C.jeremydewaardi, C.jessiehillae, C.jesusugaldei, C.jimlewisi, C.jimmilleri, C.jimwhitfieldi, C.johanvalerioi, C.johnburnsi, C.johnnoyesi, C.jorgebaltodanoi, C.jorgehernandezi, C.josealfredohernandezi, C.josefernandeztrianai, C.josehernandezcortesi, C.josemanuelperezi, C.josephinerodriguezae, C.juanmatai, C.junkoshimurae, C.kateperezae, C.luciariosae, C.luzmariaromeroae, C.manuelpereirai, C.manuelzumbadoi, C.marianopereirai, C.maribellealvarezae, C.markmetzi, C.markshawi, C.martajimenezae, C.mayrabonillae, C.meganmiltonae, C.melaniamunozae, C.michaelstroudi, C.michellevanderbankae, C.mingfangi, C.minorcarmonai, C.monikaspringerae, C.moniquegilbertae, C.motohasegawai, C.nataliaivanovae, C.nelsonzamorai, C.normwoodleyi, C.osvaldoespinozai, C.pamelacastilloae, C.paulgoldsteini, C.paulhansoni, C.paulheberti, C.petronariosae, C.ramyamanjunathae, C.randallgarciai, C.rebeccakittelae, C.robertoespinozai, C.robertofernandezi, C.rocioecheverriae, C.rodrigogamezi, C.ronaldzunigai, C.rosibelelizondoae, C.rostermoragai, C.ruthfrancoae, C.scottmilleri, C.scottshawi, C.sergioriosi, C.sigifredomarini, C.stevearonsoni, C.stevestroudi, C.sujeevanratnasinghami, C.sureshnaiki, C.torbjornekremi, C.yeimycedenoae, Leptodrepanaalexisae, L.erasmocoronadoi, L.felipechavarriai, L.freddyquesadai, L.gilbertfuentesi, L.manuelriosi, Phanerotomaalmasolisae, P.alvaroherrerai, P.anacordobae, P.anamariamongeae, P.andydeansi, P.angelagonzalezae, P.angelsolisi, P.barryhammeli, P.bernardoespinozai, P.calixtomoragai, P.carolinacanoae, P.christerhanssoni, P.christhompsoni, P.davesmithi, P.davidduthiei, P.dirksteinkei, P.donquickei, P.duniagarciae, P.duvalierbricenoi, P.eddysanchezi, P.eldarayae, P.eliethcantillanoae, P.jenopappi, Pseudophanerotomaalanflemingi, Ps.albanjimenezi, Ps.alejandromarini, Ps.alexsmithi, Ps.allisonbrownae, Ps.bobrobbinsi. HOMOLOBINAE: Exasticolusjennyphillipsae, E.randallgarciai, E.robertofernandezi, E.sigifredomarini, E.tomlewinsoni. HORMIINAE: Hormiusanamariamongeae, H.angelsolisi, H.anniapicadoae, H.arthurchapmani, H.barryhammeli, H.carmenretanae, H.carloswalkeri, H.cesarsuarezi, H.danbrooksi, H.eddysanchezi, H.erikframstadi, H.georgedavisi, H.grettelvegae, H.gustavoinduni, H.hartmanguidoi, H.hectoraritai, H.hesiquiobenitezi, H.irenecanasae, H.isidrochaconi, H.jaygallegosi, H.jimbeachi, H.jimlewisi, H.joelcracrafti, H.johanvalerioi, H.johnburleyi, H.joncoddingtoni, H.jorgecarvajali, H.juanmatai, H.manuelzumbadoi, H.mercedesfosterae, H.modonnellyae, H.nelsonzamorai, H.pamelacastilloae, H.raycypessi, H.ritacolwellae, H.robcolwelli, H.rogerblancosegurai, H.ronaldzunigai, H.russchapmani, H.virginiaferrisae, H.warrenbrighami, H.willsflowersi. ICHNEUTINAE: Oligoneuruskriskrishtalkai, O.jorgejimenezi, Paroligoneuruselainehoaglandae, P.julianhumphriesi, P.mikeiviei. MACROCENTRINAE: Austrozelejorgecampabadali, A.jorgesoberoni, Dolichozelegravitarsis (Muesebeck, 1938), D.josefernandeztrianai, D.josephinerodriguezae, Hymenochaoniakalevikulli, H.kateperezae, H.katherinebaillieae, H.katherineellisonae, H.katyvandusenae, H.kazumifukunagae, H.keithlangdoni, H.keithwillmotti, H.kenjinishidai, H.kimberleysheldonae, H.krisnorvigae, H.lilianamadrigalae, H.lizlangleyae, Macrocentrusfredsingeri, M.geoffbarnardi, M.gregburtoni, M.gretchendailyae, M.grettelvegae, M.gustavogutierrezi, M.hannahjamesae, M.harisridhari, M.hillaryrosnerae, M.hiroshikidonoi, M.iangauldi, M.jennyphillipsae, M.jesseausubeli, M.jessemaysharkae, M.jimwhitfieldi, M.johnbrowni, M.johnburnsi, M.jonathanfranzeni, M.jonathanrosenbergi, M.jorgebaltodanoi, M.lucianocapelli. ORGILINAE: Orgilusamyrossmanae, O.carrolyoonae, O.christhompsoni, O.christinemcmahonae, O.dianalipscombae, O.ebbenielsoni, O.elizabethpennisiae, O.evertlindquisti, O.genestoermeri, O.jamesriegeri, O.jeanmillerae, O.jeffmilleri, O.jerrypowelli, O.jimtiedjei, O.johnlundbergi, O.johnpipolyi, O.jorgellorentei, O.larryspearsi, O.marlinricei, O.mellissaespinozae, O.mikesmithi, O.normplatnicki, O.peterrauchi, O.richardprimacki, O.sandraberriosae, O.sarahmirandae, O.scottmilleri, O.scottmorii, Stantoniabillalleni, S.brookejarvisae, S.donwilsoni, S.erikabjorstromae, S.garywolfi, S.henrikekmani, S.luismirandai, S.miriamzunzae, S.quentinwheeleri, S.robinkazmierae, S.ruthtifferae. PROTEROPINAE: Hebichneutestricolor Sharkey & Wharton, 1994, Proteropsiangauldi, P.vickifunkae, Michenercharlesi. RHYSIPOLINAE: Pseudorhysipolisluisfonsecai, P. mailyngonzalezaeRhysipolisjulioquirosi. ROGADINAE: Aleiodesadrianaradulovae, A.adrianforsythi, A.agnespeelleae, A.alaneaglei, A.alanflemingi, A.alanhalevii, A.alejandromasisi, A.alessandracallejae, A.alexsmithi, A.alfonsopescadori, A.alisundermieri, A.almasolisae, A.alvarougaldei, A.alvaroumanai, A.angelsolisi, A.annhowdenae, A.bobandersoni, A.carolinagodoyae, A.charlieobrieni, A.davefurthi, A.donwhiteheadi, A.doylemckeyi, A.frankhovorei, A.henryhowdeni, A.inga Shimbori & Shaw, 2020, A.johnchemsaki, A.johnkingsolveri, A.gonodontovorus Shimbori & Shaw, 2020, A.manuelzumbadoi, A.mayrabonillae, A.michelledsouzae, A.mikeiviei, A.normwoodleyi, A.pammitchellae, A.pauljohnsoni, A.rosewarnerae, A.steveashei, A.terryerwini, A.willsflowersi, Bioalfapedroleoni, B.alvarougaldei, B.rodrigogamezi, Choreborogasandydeansi, C.eladiocastroi, C.felipechavarriai, C.frankjoycei, Clinocentrusandywarreni, Cl.angelsolisi, Cystomastaxalexhausmanni, Cy.angelagonzalezae, Cy.ayaigarashiae, Hermosomastaxclavifemorus Quicke sp. nov., Heterogamusdonstonei, Pseudoyeliconesbernsweeneyi, Stiropiusbencrairi, S.berndkerni, S.edgargutierrezi, S.edwilsoni, S.ehakernae, Triraphisbillfreelandi, T.billmclarneyi, T.billripplei, T.bobandersoni, T.bobrobbinsi, T.bradzlotnicki, T.brianbrowni, T.brianlaueri, T.briannestjacquesae, T.camilocamargoi, T.carlosherrerai, T.carolinepalmerae, T.charlesmorrisi, T.chigiybinellae, T.christerhanssoni, T.christhompsoni, T.conniebarlowae, T.craigsimonsi, T.defectus Valerio, 2015, T.danielhubi, T.davidduthiei, T.davidwahli, T.federicomatarritai, T.ferrisjabri, T.mariobozai, T.martindohrni, T.matssegnestami, T.mehrdadhajibabaei, T.ollieflinti, T.tildalauerae, Yeliconesdirksteinkei, Y.markmetzi, Y.monserrathvargasae, Y.tricolor Quicke, 1996. Y.woldai Quicke, 1996. The following new combinations are proposed: Neothlipsissmithi (Ashmead), new combination for Microdussmithi Ashmead, 1894; Neothlipsispygmaeus (Enderlein), new combination for Microduspygmaeus Enderlein, 1920; Neothlipsisunicinctus (Ashmead), new combination for Microdusunicinctus Ashmead, 1894; Therophilusanomalus (Bortoni and Penteado-Dias) new combination for Plesiocoelusanomalus Bortoni and Penteado-Dias, 2015; Aerophilusareolatus (Bortoni and Penteado-Dias) new combination for Plesiocoelusareolatus Bortoni and Penteado-Dias, 2015; Pneumagathiserythrogastra (Cameron) new combination for Agathiserythrogastra Cameron, 1905. Dolichozelecitreitarsis (Enderlein), new combination for Paniscozelecitreitarsis Enderlein, 1920. Dolichozelefuscivertex (Enderlein) new combination for Paniscozelefuscivertex Enderlein, 1920. Finally, Bassusbrooksi Sharkey, 1998 is synonymized with Agathiserythrogastra Cameron, 1905; Paniscozelegriseipes Enderlein, 1920 is synonymized with Dolichozelekoebelei Viereck, 1911; Paniscozelecarinifrons Enderlein, 1920 is synonymized with Dolichozelefuscivertex (Enderlein, 1920); and Paniscozelenigricauda Enderlein,1920 is synonymized with Dolichozelequaestor (Fabricius, 1804). (originally described as Ophionquaestor Fabricius, 1804).

Revision of Telothyria van der Wulp (Diptera: Tachinidae) and twenty-five new species from Area de Conservación Guanacaste in northwestern Costa Rica with a key to Mesoamerican species.

BACKGROUND: We describe 25 new species in the genus Telothyria van der Wulp, 1890 from Area de Conservación Guanacaste (ACG) in northwestern Costa Rica. All species herein described were reared from an ongoing inventory of wild-caught caterpillars spanning two families (Lepidoptera: Crambidae, and Tortricidae). Our study provides a concise description of each new species using morphology, life history, molecular data, and photographic documentation; a redescription of the genus, and its type species as well as a revised key to species of Telothyria occurring in the Mesoamerican region. We also suggest seven new synonymies resulting in 11 new combinations. NEW INFORMATION: The following 25 new species of Telothyria are described: T. aidani sp. n., T. alexanderi sp. n., T. auranticrus sp. n., T. auriolus sp. n., T. bicuspidata sp. n., T. carolinacanoae sp. n., T. clavata sp. n., T. cristata sp. n., T. diniamartinezae sp. n., T. duniagarciae sp. n., T. duvalierbricenoi sp. n., T. eldaarayae sp. n., T. erythropyga sp. n., T. fimbriata sp. n., T. fulgida sp. n., T. gloriashihezarae sp. n., T. grisea sp. n., T. harryramirezi sp. n., T. incisa sp. n., T. manuelpereirai sp. n., T. obscura sp. n., T. omissa sp. n., T. osvaldoespinozai sp. n., T. peltata sp. n., and T. ricardocaleroi sp. n.The following are proposed by Fleming & Wood as new generic synonyms of Telothyria: Comatacta Coquillett Syn. n., Floradalia Thompson Syn. n., Ptilomyia Curran Syn. n., Ptilomyiopsis Townsend Syn. n., Ptilomyoides Curran Syn. n., Euptilomyia Syn. n., Eutelothyria Townsend Syn. n.The following new combinations are proposed as a result of the new synonymies: Telothyria bequaerti (Curran, 1925) Comb. n., Telothyria cruenta (Giglio-Tos, 1893) Comb. n., Telothyria frontalis (Townsend, 1939) Comb. n., Telothyria insularis (Curran, 1927) Comb. n., Telothyria itaquaquecetubae (Townsend, 1931) Comb. n., Telothyria major (Thompson, 1963) Comb. n., Telothyria micropalpus (Curran, 1925) Comb. n., Telothyria minor (Thompson, 1963) Comb. n., Telothyria nautlana (Townsend, 1908) Comb. n., Telothyria plumata (Curran, 1925) Comb. n., Telothyria trinitatis (Thompson, 1963) Comb. n., Telothyria variegata (Fabricius, 1805) Comb. n. Musca tricincta Fabricius is synonymized under Telothyria variegata Fabricius, Syn. n. Telothyria schineri Fleming & Wood nom. n. is proposed as a replacement name for Miltogramma brevipennis Schiner.Additionally we provide redescriptions of two previously named species: the type species Telothyria cupreiventris (van der Wulp) due to its being the type species, and Telothyria relicta (van der Wulp) due to its having been reared as an outcome of the inventory.

A new genus and new species in the tribe Uramyini (Diptera: Tachinidae) from Area de Conservación Guanacaste in northwestern Costa Rica.

BACKGROUND: We describe one new genus and its one new species from Area de Conservación Guanacaste (ACG) in northwestern Costa Rica. Our study provides a concise description of this new species using morphology, life history, molecular data and photographic documentation. NEW INFORMATION: Chorotegamyia gen. n. is described, along with its type species, Chorotegamyia aureofacies sp. n. A modified key to the Uramyini is given to further elucidate the tribe.