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Zootaxa 3858 (1): 001–090 www.mapress.com /zootaxa / Copyright © 2014 Magnolia Press ISSN 1175-5326 (print edition) Monograph ZOOTAXA ISSN 1175-5334 (online edition) http://dx.doi.org/10.11646/zootaxa.3858.1.1 http://zoobank.org/urn:lsid:zoobank.org:pub:D1CCF02B-4314-4537-A64F-0372715E3F93 ZOOTAXA 3858 Revision of the New World species of Houghia Coquillett (Diptera, Tachinidae) reared from caterpillars in Area de Conservación Guanacaste, Costa Rica ALAN J. FLEMING1,4, D. MONTY WOOD1, M. ALEX SMITH2, WINNIE HALLWACHS3 & DANIEL H. JANZEN3 1 Canadian National Collection of Insects, Agriculture and Agri-Food Canada, 960 Carling Avenue, Ottawa, ON, Canada, K1A 0C6 2 Department of Integrative Biology & the Biodiversity Institute of Ontario, University of Guelph, Guelph, ON, Canada, N1G 2W1 3 Department of Biology, University of Pennsylvania, Philadelphia, PA 19104, USA 4 Corresponding author. E-mail: [email protected] Magnolia Press Auckland, New Zealand Accepted by J. OHara: 24 Jul. 2014; published: 2 Sept. 2014 ALAN J. FLEMING, D. MONTY WOOD, M. ALEX SMITH, WINNIE HALLWACHS & DANIEL H. JANZEN Revision of the New World species of Houghia Coquillett (Diptera, Tachinidae) reared from caterpillars in Area de Conservación Guanacaste, Costa Rica (Zootaxa 3858) 90 pp.; 30 cm. 2 Sept. 2014 ISBN 978-1-77557-481-1 (paperback) ISBN 978-1-77557-482-8 (Online edition) FIRST PUBLISHED IN 2014 BY Magnolia Press P.O. Box 41-383 Auckland 1346 New Zealand e-mail: [email protected] http://www.mapress.com/zootaxa/ © 2014 Magnolia Press All rights reserved. No part of this publication may be reproduced, stored, transmitted or disseminated, in any form, or by any means, without prior written permission from the publisher, to whom all requests to reproduce copyright material should be directed in writing. This authorization does not extend to any other kind of copying, by any means, in any form, and for any purpose other than private research use. ISSN 1175-5326 (Print edition) ISSN 1175-5334 (Online edition) 2 · Zootaxa 3858 (1) © 2014 Magnolia Press FLEMING ET AL. Table of contents Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Materials and methods. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 DNA Barcoding results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 Conspectus of the genus Houghia Coquillett . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 Generic Synonyms of Houghia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 Previously described species included in Houghia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 Lectotype designations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 Diagnosis of the Genus Houghia. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 Key to Males of Houghia Reared from Caterpillars in ACG, Northwestern Costa Rica . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30 Species diagnoses and descriptions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32 Houghia aerata Fleming & Wood, sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32 Houghia aurifera Fleming & Wood, sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34 Houghia biseriata Fleming & Wood, sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35 Houghia bivittata Fleming & Wood, sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37 Houghia blancoi Fleming & Wood, sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39 Houghia brevipilosa Fleming & Wood, sp. nov.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40 Houghia chavarriae Fleming & Wood, sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42 Houghia confinis Fleming & Wood, sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43 Houghia crypta (Townsend, 1934 402) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45 Houghia delospilota Fleming & Wood, sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47 Houghia destituta Fleming & Wood, sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48 Houghia fimbriata Fleming & Wood, sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50 Houghia gracilis Fleming & Wood, sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51 Houghia graciloides Fleming & Wood, sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53 Houghia griseifrons Fleming & Wood, sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55 Houghia inflatipalpis Fleming & Wood, sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56 Houghia latigena Fleming & Wood, sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58 Houghia latilobus Fleming & Wood sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59 Houghia longicercus Fleming & Wood, sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61 Houghia longipilosa Fleming & Wood, sp. nov.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62 Houghia luteiventris Fleming & Wood sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64 Houghia macilenta Fleming & Wood, sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66 Houghia marini Fleming & Wood, sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67 Houghia matarritai Fleming & Wood, sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69 Houghia nigrofemur Fleming & Wood, sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71 Houghia ochrofemur Fleming & Wood, sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72 Houghia omissa Fleming & Wood, sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 74 Houghia pallida Fleming & Wood, sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75 Houghia parmata Fleming & Wood, sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77 Houghia pilosifrons Fleming & Wood, sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78 Houghia romeroae Fleming & Wood, sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 80 Houghia sexmaculata Fleming & Wood, sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81 Houghia spathulata Fleming & Wood, sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82 Houghia triangularis Fleming and Wood, sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84 Houghia velutina Fleming & Wood sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85 Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87 Literature cited. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87 HOUGHIA OF ACG, COSTA RICA Zootaxa 3858 (1) © 2014 Magnolia Press · 3 Abstract Thirty-five species of the genus Houghia Coquillett (Tachinidae, Exoristinae, Goniini) are described, 34 new and one previously described, all reared from various species of caterpillars collected in Area de Conservación Guanacaste (ACG), northwestern Costa Rica. A matrix of character states and a key for the identification of the species are also provided. By coupling morphology, life history and molecular data, with photographic documentation, a clear and concise description of each species is provided. The following 34 new species of Houghia are described, all authored by Fleming and Wood: H. aerata sp. nov., H. aurifera sp. nov., H. biseriata sp. nov., H. bivittata sp. nov., H. blancoi sp. nov., H. brevipilosa sp. nov., H. chavarriae sp. nov., H. confinis sp. nov., H. delospilota sp. nov., H. destituta sp. nov., H. fimbriata sp. nov., H. gracilis sp. nov., H. graciloides sp. nov., H. griseifrons sp. nov., H. inflatipalpis sp. nov., H. latigena sp. nov., H. latilobus sp. nov., H. longicercus sp. nov., H. longipilosa sp. nov., H. luteiventris sp. nov., H. macilenta sp. nov., H. marini sp. nov., H. matarritai sp. nov., H. nigrofemur sp. nov., H. ochrofemur sp. nov., H. omissa sp. nov., H. pallida sp. nov., H. parmata sp. nov., H. pilosifrons sp. nov., H. romeroae sp. nov., H. sexmaculata sp. nov., H. spathulata sp. nov., H. triangularis sp. nov., and H. velutina sp. nov. The following are proposed by Wood as new synonyms of Houghia: Actinoprosopa Townsend syn. nov., Agrarialia Curran syn. nov., Anhangabahuia Townsend syn. nov., Aridalia Curran syn. nov., Bolohoughia Townsend syn. nov., Carceliocephala Townsend syn. nov., Chrysohoughia Townsend syn. nov., Eumacrohoughia Townsend syn. nov., Macrohoughia Townsend syn. nov., Orohoughia Townsend syn. nov., Pammaerus Aldrich syn. nov., Pararrhinactia Townsend syn. nov., Petrargyrops Townsend syn. nov., Sisyrohoughia Townsend syn. nov., Tapajohoughia Townsend syn. nov., and Verrugomyia Townsend syn. nov. New combinations are proposed by Wood as a result of the new synonymies, as follows: H. analis (Townsend) comb. nov., H. approximata (van der Wulp) comb. nov., H. aurata (Townsend) comb. nov., H. aurometallica (Townsend) comb. nov., H. bistrigata (van der Wulp) comb. nov., H. calcarata (van der Wulp) comb. nov., H. chlorescens (Townsend) comb. nov., H. crypta (Townsend) comb. nov., H. facialis (Townsend) comb. nov., H. impedita (van der Wulp) comb. nov., H. lateralis (Curran) comb. nov., H. leptotrichopa (Brauer & Bergenstamm) comb. nov., H. maris (Townsend) comb. nov., H. marmorata (Townsend) comb. nov., H. minor (Thompson) comb. nov., H. nuda (Townsend) comb. nov., H. orbitalis (Townsend) comb. nov., H. orbitalis (Curran) comb. nov., H. parva (Townsend) comb. nov., H. plagioides (van der Wulp) comb. nov., H. punctiger (Townsend) comb. nov., H. quadra (Wiedemann) comb. nov., H. sexualis (Curran) comb. nov., H. similis (Townsend) comb. nov., H. simillima (Thompson) comb. nov., H. sordida (van der Wulp) comb. nov., H. tenuiseta (Macquart) comb. nov., and H. tropica (Townsend) comb. nov. Houghia orbitalis (Curran, 1934a, described in Sturmia), junior homonym of H. orbitalis (Townsend, 1927, described in Verrugomyia) is synonymized by Wood with Houghia leptotrichopa (Brauer & Bergenstamm, 1891), and need not be renamed while in synonymy. Lectotypes are designated by Wood for the following ten valid species of Houghia: Anisia approximata van der Wulp, 1890, Actinoprosopa facialis Townsend, 1927, Masicera impedita van der Wulp, 1890, Macrohoughia marmorata Townsend, 1927, Eumacrohoughia minor Thompson, 1963, Prospherysa plagioides van der Wulp, 1890, Petrargyrops punctiger Townsend, 1927, Carceliocephala simillima Thompson, 1963, Masicera sordida van der Wulp, 1890, and Masicera tenuiseta Macquart, 1846: 292. Key words: Houghia, Diptera, Tachinidae, Exoristinae, Goniini, tropical rain forest, tropical dry forest, parasitoid fly, host-specificity, caterpillars Introduction Area de Conservación Guanacaste (ACG) is a model of conservation effort through biodiversity development. As a result of the tireless efforts of its staff and collaborators, more than half a million wild-caught lepidopteran larvae have been reared for their parasitoids (Janzen et al. 2009, Janzen & Hallwachs 2011, Fernandez-Triana et al. 2014). This provides an unprecedented amount of data, providing invaluable information on parasitoid biology and associated hosts. The most speciose fly parasitoids encountered belong to the family Tachinidae. The Neotropical Region has a tachinid fauna vastly more diverse than anything indicated in any publication. Although the number of named species catalogued by Guimarães (1971), some 2864 species, is larger than that of any other faunal region, this is undoubtedly a small fraction of what exists in nature, based on what has recently been discovered in Costa Rica, and what is already present in other collections. The most speciose elements of the fauna seem to occur in the upper elevations and cloud forests that extend from the western slopes of the Sierra Madre Occidentale in Mexico to both slopes of the Andes from Colombia south to Bolivia, and numerous species of Houghia have been found throughout this mountain chain. The New World genus Houghia was erected by Coquillett (1897) for two specimens collected at Tifton, 4 · Zootaxa 3858 (1) © 2014 Magnolia Press FLEMING ET AL. Georgia, USA, by G.R. Pilate, which he described as Houghia setipennis Coqullett. The genus remained with this single species until Reinhard (1967) added a second species, H. nigripalpis Reinhard. Wood (1987) added five more described Nearctic species, under the name Eumasicera Townsend with Houghia in synonymy, not realizing that Houghia is the senior synonym (O’Hara & Wood 1998). By adding 29 more previously named species that were described in nearly as many genera, this paper is the first to substantially enlarge the genus (under Conspectus, below). Character states that in combination provide a diagnosis of Houghia are also presented below. Coquillett published his two specimens of H. setipennis as females, but they are in fact males. This simple error introduced one of the most important characters of the genus, an error often made by subsequent authors: males of all species of Houghia, including H. setipennis, have two pairs of proclinate orbital setae that are as well developed as those in females. This female-like character state is not present in males of similar-looking genera such as Carcelia Robineau-Desvoidy (Eryciini) and Hyphantrophaga Townsend (Goniini). Correspondingly, the width of the frons is only slightly different in the two sexes. Unfortunately, males belonging to many other tachinid genera with a broad frons also have two pairs of proclinate orbital setae and male Houghia need to be distinguished from these other genera by other character states, as elaborated below under “Character states of the genus Houghia”. As in all members of the tribe Goniini, females deposit microtype eggs on foliage that are then swallowed by the host. However, in one species described by Thompson (1963: 402) under the name Carceliocephala simillima Thompson, these eggs are atypical, resembling a bivalve rather than a limpet, and their structure is not understood because of a paucity of material. Unfortunately this lack of a single distinctive generic feature makes females even more difficult to tell apart than males, but this is generally true among most genera of tachinids. Wood (1987) suggested that the lack of differentiation of frontal width was associated with female-finding behaviour of continually visiting prospective sites on the wing in search of waiting females. This contrasts with waiting at a particular location (“aggregation site”) for a female to arrive and then competing with other waiting males for her attention, as do most tachinids. Continual searching is presumably more energy intensive than is waiting but avoids the male/male competition that is usually present at aggregation sites. Continual searching may also be an effective strategy in localities with massive amounts of vegetation and hilltops draped in cold clouds, or where females occur at very low density. Houghia is not limited to any one biome within ACG, and can be found parasitizing a variety of lepidopteran hosts throughout the research area. As part of the tribe Goniini, Houghia females lay their microtype eggs by the hundreds on foliage, the egg is then swallowed with its leaf fragment when the caterpillar is feeding, and the newly hatched maggot finds its way to a safe part of the caterpillar body, where it lives until it continues growth and consumption of the caterpillar. Only very rarely does this maggot not kill the caterpillar (e.g., DeVries (1983) once reared an adult butterfly from a caterpillar that had already produced viable tachinid larvae), but the ACG inventory has had this happen only once out of 25,000+ tachinid rearings. In this paper, 34 of the 35 species of Houghia reared from caterpillars collected in ACG are described as new, based on differences in external morphology, male terminalia, and CO1 (cox1 or cytochrome oxidase 1) gene sequences, and on comparison by DMW with other species of Houghia from other places. However, this is by no means an indication of the total number of species of Houghia even in such a small country as Costa Rica. The Houghia collection at the Instituto Nacional de Biodiversidad, in Santo Domingo de Heredia, Costa Rica, consists of many species. The majority of specimens belong to only two species, and although both are common in ACG and elsewhere in Guanacaste Province, neither has been reared. The remaining male specimens in the INBio collection appear to be species different from those reared in ACG. Some may belong among the 34 described below but either were not successfully sequenced, were in poor condition, or did not look sufficiently similar to any of the 34 to be included as paratypes, suggesting that the Houghia fauna in the rest of the country is quite different from that of ACG (as is the climate) and requires much additional study. There may also be a small number of apparent species of Houghia that have been reared by the ACG inventory that at present can be distinguished only by their gene sequences (henceforth referred to as DNA barcodes) and host records; AJF and DMW have elected to leave them for later description when additional material is available. HOUGHIA OF ACG, COSTA RICA Zootaxa 3858 (1) © 2014 Magnolia Press · 5 Materials and methods Acronyms for Depositories AMNH BMNH CNC MZSP NHMW SENASA USNM ZMAM INBio American Museum of Natural History, New York, USA The Natural History Museum, London, United Kingdom Canadian National Collection of Insects, Arachnids and Nematodes, Ottawa, Canada Musei de Zoologia Universidade de São Paulo, São Paulo, Brazil Naturhistorisches Museum Wien, Vienna, Austria Servicio Nacional de Sanidad Agraria, La Molina, Lima, Peru National Museum of Natural History, Washington, DC, USA Institute of Systematics and Biology of Populations, Zoological Museum, University of Amsterdam, The Netherlands. Instituto Nacional de Biodiversidad, Santo Domingo de Heredia, Costa Rica Geographic Area of the Study and Rearing Intensity. All flies and rearing information described here were found by the 35+ year-old ongoing inventory of the caterpillars, their food plants and their parasitoids of the dry forest, rain forest, cloud forest, and intergrades, in the 125,000+ ha terrestrial portion of ACG (Rodriguez et al. 2013, 2014, Janzen et al. 2009, Janzen and Hallwachs 2011, Smith et al. 2006, 2007, 2008, 2012). The tachinid rearing methods are described at http://janzen.bio.upenn.edu/caterpillars/methodology/how/parasitoid_husbandry.htm. This inventory has reared 600,000+ wild-caught ACG caterpillars since 1978. All frequencies of parasitization reported here need to be considered against this background inventory. Dissections. Adult fly terminalic dissections followed standard practice (O’Hara 1983, 2002). Due to the destructive nature of the dissections, specimens to be dissected were selected based on their inferior photogenic quality but certain identity based on gene sequences in common with their respective holotypes. The abdomen was gently pried off by applying gentle pressure to the underside until a slight snap was felt. If the abdomen did not fall off at this point it was gently rocked from side to side until it could be removed from the thorax. The abdomen was placed in a small crucible of 10% NaOH placed on a hotplate to heat at sub-boiling temperature for 15 minutes. With maceration complete, the abdomen was then placed in a dissection plate with 50% acetic acid. The submerged abdomen was held in place with a bent-tipped probe. The terminalia were carefully cut away from the abdomen, using a wooden probe armed with a sharpened minuten pin. Under 16X magnification the genital capsule was dissected out with this tool. After a further 15 minutes in acetic acid, the entire genital capsule was then placed in a microvial containing glycerin, and attached to the specimen pin for storage. Microvials are used instead of slide mounting because of the 3-D form of the terminalia. The dissected abdomen was then bathed in a series of dehydration steps; it was passed through ethanol baths (50%, 95%) and then into acetone. The dehydrated and dry abdomen was then reattached to the specimen using shellac glue. Terminology used for terminalia (which refers here only to the sclerotized parts of the genitalia, and not to the internal structures, although often refered to as genitalia) and other body parts follows Cumming and Wood (2009). Imaging. Habitus photographs were taken using a Canon 40D digital SLR, using a 65mm Macro Photo Lens 1:2.8 (MP-E 65mm), double mounted on a Cognisys Inc. Stackshot tripod electronically controlled focus stacking rail system on a Kaiser (RS 1 with RA-1 arm) copy stand. Images were shot in aperture priority, allowing the camera to control shutter speed at f/6.3, over 18 images at equal distance increments. Illumination was with a reflective dome placed over a 110 LED ring light. Photographs of male terminalia were taken using a Canon S110 digital camera adaptor mounted to the eyepiece of a Leitz-Wetzlar dissecting scope. Preparations were mounted on a depression slide in a small quantity of Rexall brand hand sanitizer gel (NPN# 80007138). Hand sanitizer gel provided a more viscous mounting medium than did glycerin traditionally used for the same purpose. This allowed the specimen to remain steady in a given position, and therefore provided a better environment for photography. After mounting and photographing, the terminalia were rinsed in a small quantity of pure distilled water, before being replaced in the glycerin-filled microvial. Photographic series were then prepared using Photoshop CS6, and Zerene Stacker Software v1.04. In order to maximize quality and depth of field, photos were digitally stacked to produce the final composite image. Wherever a specimen label has been examined, the information is presented using the following symbols: /, 6 · Zootaxa 3858 (1) © 2014 Magnolia Press FLEMING ET AL. indicates the end of a line; //, indicates the end of a label. Labels are presented from top (closest to the specimen) to bottom, with any comments on the label being given in square brackets. Voucher specimen management. For the purposes of this study approximately 2000 specimens reared from ACG and belonging to the genus Houghia were examined. Most relevant for this taxonomic paper is that each reared caterpillar receives a unique voucher code in the format of yy-SRNP-xxxxx, the record of which is available at http://janzen.bio.upenn.edu/caterpillars/database.lasso. Any parasitoid emerging from this caterpillar receives the same voucher code, such that if multiple parasite specimens emerge from one host they all bear the same number representing one rearing event. Then if/when later it is dealt with individually, each parasite receives a second voucher code unique to it, in the format of DHJPARxxxxxxx. This voucher code may likewise be sought at http://janzen.bio.upenn.edu/caterpillars/database.lasso. To date, all DHJPARxxxxxxx coded tachinids have had a leg removed for attempted DNA barcoding at the Biodiversity Institute of Ontario (BIO) in Guelph, Ontario, Canada, with all collateral data and all successful barcodes permanently and publically deposited in the Barcode of Life Data System (BOLD, www.boldsystems.org) (Ratnasingham & Hebert 2007), and later migrated to GenBank as well. A neighbour-joining (NJ) tree (Saitou & Nei, 1987) tree for all Houghia reared and DNA barcoded by this inventory to date is included as Supplemental Appendix 1. As the inventory is continually growing, new specimens and species can be found by searching the genus Houghia in BOLD. Each barcoded specimen also has an accession code from the Barcode of Life Data System (BOLD) and GenBank. All Tachinidae reared by this ACG inventory (except for the genus Belvosia Robineau-Desvoidy), are permanently deposited in the CNC, except for those that were destroyed or donated to other collections as paratypes or supporting material. Formally, all inventory Tachinidae have been collected under Costa Rican government research permits issued to DHJ, and likewise exported, under permit by DHJ from Costa Rica to Philadelphia, and then to Ottawa. All tachinid identification by the inventory has been done by DHJ in coordination with a) combination of visual inspection by AJF (for Houghia and other genera under revision) and DMW, b) DNA barcoding by authors BIO and MAS, and by BOLD, and c) correlation with host caterpillar identifications by DHJ and WH through the inventory itself. Dates of capture of each holotype are the dates of eclosion of the fly, and not the date of capture of the caterpillar, since the fly eclosion date is much more representative of the time when that fly species is on the wing, than is the time of capture of the caterpillar. However, the collector listed is the parataxonomist who found the caterpillar, rather than the person who retrieved the newly eclosed fly and processed it by freezing, pinning, labeling and oven-drying. Fly biology and parasitization by these flies will be the subject of later papers. DNA Barcoding. DNA extracts were prepared from single legs using a glass fibre protocol (Ivanova et al. 2006). Extracts were re-suspended in 30 μl of dH2O, and a 658 bp region near the 5’ terminus of the CO1 gene was amplified using standard primers following established protocols (Smith et al. 2006, 2007, 2008). If this 658 bp amplification was not successful, smaller overlapping sequences were targeted using the same primer pair as above combined with the internal primers. The primer information for individual sequences can be retrieved from the Barcode of Life Data System (BOLD) (Ratnasingham & Hebert 2007). Primer sequence information is detailed in Table 1. DNA Barcoding results CO1 fragments generated were from amplifications where agarose gels were free of double bands and the resultant sequences were free of systematic heteroplasmy – both of which can be signs of the inadvertent amplification of pseudogenes (Bensasson et al. 2001). The DNA barcodes were AT biased (average GC content 29.64%) which is characterstic of insect mitochondrial DNA (Zhang & Hewitt 1997). The variation within a species (mean pairwise distance 0.176%, max pairwise distance, 1.55%) was much lower than the variation between species (mean pairwise distances 8.38%). A CO1 sequence of some length was generated from 927 of 954 specimens extracted. Unless otherwise noted (Houghia pallida DHJPAR0008432 496 bp, Houghia gracilis, DHJPAR0008435 598 bp, Houghia longicercus DHJPAR0008473 574 bp, Houghia delospilota DHJPAR0008837 502 bp), the analyzed sequences were longer than 600 bp and free of ambiguities. All sequence data for the Houghia specimens examined can be accessed on BOLD using the public DOI: dx.doi.org/10.5883/DS-ASHOU. Accessions and collection information (BOLD, and GenBank) for all sequences can be viewed in Supplemental Appendix 2. Sequences were HOUGHIA OF ACG, COSTA RICA Zootaxa 3858 (1) © 2014 Magnolia Press · 7 then examined and used as a preliminary sorting tool providing “buckets” into which each species could be divided, these buckets were then confirmed with morphological and species biology data. By using all three data sets we provide a solid concept of each species, based on CO1 barcodes, as well as the more traditional morphological determinations. TABLE 1. Primers used to amplify the DNA barcode region of COI. The prefix “C_” denotes a cocktail of two primers and the suffix “_t” denotes a primer with a M13 tail for sequencing. Primer Name Contig (C_) Primer Sequence ( 5' to 3') Reference C LepFolF LepF1:LCO1490 LCO1490 GGTCAACAAATCATAAAGATATTGG Folmer et al. (1994) LepF1 ATTCAACCAATCATAAAGATATTGG Hebert et al. (2004) MLepF1 GCTTTCCCACGAATAAATAATA Hajibabaei et al. (2005) RonMWASPdeg_t1 TGTAAAACGACGGCCAGTGGWTCWCCW GATATAKCWTTTCC Smith et al. (2009) LCO1490_t1 TGTAAAACGACGGCCAGTGGTCAACAAA TCATAAAGATATTGG LCO1490 with M13 tail MLepR2 GTTCAWCCWGTWCCWYCCATTTTC unpublished MLepR1 CCTGTTCCAGCTCCATTTTC Hajibabaei et al. (2006) LepR1 CTCCWCCAGCAGGATCAAAA Hebert et al. (2004) C_ANTMR1D RonIIdeg_R:AMR 1deg_R Smith et al. (2005) AMR1deg_R CAWCCWGTWCCKRMNCCWKCAT Smith et al. (2005) RonIIdeg_R GGRGGRTARAYAGTTCATCCWGTWCC Simon et al. (1994) C LepFolR LepR1:HCO2198 HCO2198 tl CAGGAAACAGCTATGACTAAACTTCGGGT HCO2198 with M13 tail GACCAAAAAATCA HCO2198 TAAACTTCAGGGTGACCAAAAAATCA Folmer et al. (1994) Conspectus of the genus Houghia Coquillett The study of New World tachinids by DMW has been ongoing for nearly 50 years. Before it began, each species tended to be in its own genus. For example, the genus Houghia originally embraced a single species, H. setipennis Coquillett. The other species here included in Houghia were first placed in catch-all genera such as Tachina Meigen, Masicera Macquart, Phorocera Robineau-Desvoidy, Prospherysa van der Wulp, Anisia van der Wulp, Sisyropa Brauer & Bergenstamm, Sturmia Robineau-Desvoidy, and Hypostena Meigen. These genera are now regarded as belonging to other tribes and subfamilies in the Tachinidae. Townsend eventually proposed 14 genera for the species listed below, dispersed among several tribes, the majority in Trypherini. Townsend started his taxonomic career with a better understanding of tachinid generic characteristics than had most of his predecessors, but he was strongly influenced, as were all of his contemporaries, by the use of chaetotaxy as a means of defining the genera of these flies. However, as the years passed he became more and more obsessed with the need for splitting genera into smaller and smaller units, many of them based on character states that are barely helpful for separating species, and were in some cases figments of his imagination. We now know that he had before him only a small fraction of the extant species that show every imaginable combination and recombination of his character states. He also must have known, but chose to ignore, the fact that some character states, especially of the head and abdomen, varied markedly between the two sexes of the same species. Townsend (1935b: 38) believed that “Two species, the progeny from whose crossing is sexually infertile, belong to separate natural genera” to which van Emden (1945: 389–390) remarked “the adoption of [this] principle implies the application of the generic unit to every unit considered to be a species in general zoological practice” (see O’Hara 2013). As the acquisition and study of New World tachinids progressed over the years it became apparent that many 8 · Zootaxa 3858 (1) © 2014 Magnolia Press FLEMING ET AL. of the species Townsend had each assigned to its own unique genus were scarcely different from one another and that females could usually not be associated confidently with males, adding to the difficulty. This was particularly true for species that have been synonymized in this paper. In the New World there are many dozens, probably hundreds, of different species that all share most of their character states with Houghia setipennis, and yet as a group they all share much the same appearance or “Gestalt”, so we could see no reason why they should not all be placed in Houghia, hence the new synonymies. Sabrosky and Arnaud (1965: 1094) were the first to begin this process, abandoning the Trypherini and introducing a new tribe Eumasicerini to include the North American species of Houghia, which was also applied by Guimarães (1971) to several, but not all, the Central and South American species of Houghia. We know of no unique character state that occurs in all members of the genus Houghia, so they must be recognized by a combination of character states that excludes the members of all other genera. When Coquillett originally proposed this genus for Houghia setipennis Coquillett, it was based on the presence of a row of setae on vein R1, a character state regarded at that time as worthy of separate generic rank. However, several other species of Houghia that were described in other genera have this character state, including Coquillett’s own Hypostena setinervis. An unnamed North American species that is otherwise similar to Houghia setipennis lacks these setae. In addition to having a characteristic facies, or “Gestalt” that is a feature of Houghia, the females lay microtype eggs (although most species have yet to be investigated for this trait), described in detail for five Trinidad species of Houghia by Thompson (1963: 323–341) under the generic names Carceliocephala Townsend, Eumacrohoughia Townsend, and Petrargyrops Townsend. The first instar larvae of some of these species have a simple curved labrum characteristic of many tachinids, but most have a labrum with a characteristic small dorsal angular projection, possibly a device for rupturing the egg shell as also occurs in other first instar larvae of microtype-egglaying species of the tribe Goniini (Thompson 1963, as part of the subfamily Goniinae; Herting 1984, as Goniini, and O’Hara and Wood 2004, as Goniini). Generic Synonyms of Houghia The new synonyms in the list below are proposed by DMW. Houghia Coquillett, 1897: 32, 118. Type species: Houghia setipennis Coquillett, 1897, by original designation. Eumasicera Townsend, 1909: 249. Type species: Eumasicera coccidella Townsend, 1909, by original designation. Synonymy by Wood (1987: 1206), as discussed by O’Hara & Wood (1998: 754, 761). Verrugomyia Townsend, 1927: 217. Type species: Verrugomyia orbitalis Townsend, 1927, by original designation. Syn. nov. Macrohoughia Townsend, 1927: 261. Type species: Macrohoughia marmorata Townsend, 1927, by original designation. Syn. nov. Bolohoughia Townsend, 1927: 275. Type species: Bolohoughia aurometallica Townsend, 1927, by original designation. Syn. nov. Petrargyrops Townsend, 1927: 275. Type species: Petrargyrops punctiger Townsend, 1927, by original designation. Syn. nov. Sisyrohoughia Townsend, 1927: 275. Type species: Sisyrohoughia similis Townsend, 1927, by original designation. Syn. nov. Eumacrohoughia Townsend, 1927: 275. Type species: Eumacrohoughia nuda Townsend, 1927, by original designation. Syn. nov. Actinoprosopa Townsend, 1927: 258. Type species: Actinoprosopa facialis Townsend, 1927, by original designation. Syn. nov. Pammaerus Aldrich, 1927: 24. Type species: Sisyropa leptotrichopa Brauer & Bergenstamm, 1891, by original designation. Syn. nov. Anhangabahuia Townsend, 1931: 470. Type species: Anhangabahuia analis Townsend, 1931, by original designation. Syn. nov. Carceliocephala Townsend, 1934: 402. Type species: Carceliocephala crypta Townsend, 1934, by original designation. Syn. nov. HOUGHIA OF ACG, COSTA RICA Zootaxa 3858 (1) © 2014 Magnolia Press · 9 Orohoughia Townsend, 1934: 403. Type species: Orohoughia aurata Townsend, 1927, by original designation. Syn. nov. Tapajohoughia Townsend, 1934: 405. Type species: Tapajohoughia tropica Townsend, 1934, by original designation. Syn. nov. Agrarialia Curran, 1934b: 469. Type species: Agrarialia sexualis Curran, 1934, by original designation. Syn. nov. Aridalia Curran, 1934b: 469. Type species: Aridalia lateralis Curran, 1934, by original designation. Syn. nov. Chrysohoughia Townsend, 1935a: 231. Type species: Chrysohoughia chlorescens Townsend, 1935, by original designation. Syn. nov. Pararrhinactia Townsend, 1935a: 233. Type species: Pararrhinactia parva Townsend, 1935, by original designation. Syn. nov. Pacidianus Reinhard, 1943: 88. Type species: Pacidianus hirsutus Reinhard, 1943 [= Eumasicera coccidella Townsend, 1909], by original designation. Synonymy with Eumasicera by Wood (1987: 1206, with Houghia in synonymy), see O’Hara & Wood (1998: 756). Previously described species included in Houghia The new combinations below are proposed by DMW and result from the new generic synonymy above and the examination of the type material of other nominal species. analis Townsend, 1931: 471 (Anhangabahuia). Holotype male (USNM) [examined by DMW]. Type locality: Brazil, São Paulo, Itaquaquecetuba. Comb. nov. approximata van der Wulp, 1890c: 202 (Anisia). Lectotype male (published as female) (BMNH), by designation herein (see Lectotype Designations section). Type locality: Mexico, Guerrero, Venta de Zopilote, 2800 feet. Comb. nov. aurata Townsend, 1934: 403 (Orohoughia). Holotype female (USNM) [examined by DMW]. Type locality: Brazil, Para, Río Tapajós, Bôa Vista. Comb. nov. aurometallica Townsend, 1927: 291 (Bolohoughia). Holotype male (USNM) [examined by DMW]. Type locality: Brazil, São Paulo, Itaquaquecetuba. Comb. nov. bistrigata van der Wulp, 1890a: 109 (Masicera). Holotype female (BMNH) [examined by DMW]. Type locality: Mexico, Tabasco, Teapa. Comb. nov. calcarata van der Wulp, 1890b: 114 (Masicera). Holotype female (BMNH) [examined by DMW]. Type locality: Mexico, Morelos, Cuernavaca. Comb. nov. chlorescens Townsend, 1935a: 231 (Chrysohoughia). Holotype female (BMNH) [examined by DMW]. Type locality: Trinidad. Comb. nov. coccidella Townsend, 1909: 249 (Eumasicera). Holotype female (USNM) [examined by DMW]. Type locality: USA, Massachusetts, Melrose Highlands. hirsuta Reinhard, 1943: 89 (Pacidianus). Holotype male (CNC) [examined by DMW]. Type locality: USA, Michigan, Agriculture College. Synonymy with coccidella established by Wood in O’Hara & Wood (1998: 771). crypta Townsend, 1934: 402 (Carceliocephala). Holotype male (USNM) [examined by DMW]. Type locality: Brazil, Para, Río Tapajós, Bôa Vista. Comb. nov. facialis Townsend, 1927: 283 (Actinoprosopa). Lectotype male (USNM) by designation herein (see Lectotype Designations section). Type locality: Brazil, São Paulo, Itaquaquecetuba. Comb. nov. impedita van der Wulp, 1890a: 107 (Masicera). Lectotype male (published as female) (BMNH), by designation herein (see Lectotype Designations section). Type locality: Mexico, Morelos, Cuernavaca. Comb. nov. lateralis Curran, 1934b: 469 (Aridalia). Lectotype female (AMNH), by fixation of Arnaud (1963: 107) (mention of “Holotype female” from Panama in AMNH is regarded as a lectotype fixation) [examined by DMW]. Type locality: Panama, Canal Zone, Corozal. Comb. nov. leptotrichopa Brauer & Bergenstamm, 1891: 347 (Sisyropa). Syntypes, two males [published as females] (NHMW) [examined by DMW]. Type locality: Brazil, Comb. nov. orbitalis Curran, 1934a: 515 (Sturmia). Holotype male (AMNH) [examined by DMW]. Type locality: Guyana 10 · Zootaxa 3858 (1) © 2014 Magnolia Press FLEMING ET AL. (published as British Guiana), Bartica, Kartabo. Syn. nov. Junior secondary homonym of Verrugomyia orbitalis Townsend, 1927. Synonymy is based on a male from Suriname in the CNC that had been compared with, and considered conspecific with, the types of both Sisyropa leptotrichopa and Sturmia orbitalis, suggesting that the latter should become a synonym of the former. However, these comparisons predate the present study by many years, and because of the large number of similar species of Houghia in South America they may not be correct. Nevertheless, the synonymy is provisionally proposed here, which makes the renaming of S. orbitalis Curran unnecessary. maris Townsend, 1929: 374 (Actinoprosopa). Holotype male (USNM) [examined by DMW]. Type locality: Brazil, São Paulo, Itaquaquecetuba. Comb. nov. marmorata Townsend, 1927: 325 (Macrohoughia). Lectotype male (USNM) by designation herein (see Lectotype Designations section). Type locality: Brazil, São Paulo, Itaquaquecetuba. Comb. nov. minor Thompson, 1963: 333 (Eumacrohoughia). Lectotype male (CNC) by designation herein (see Lectotype Designations section). Type locality: Trinidad, [village of] Brazil. Comb. nov. nigripalpis Reinhard, 1967: 108 (Houghia). Holotype male (CNC) [examined by DMW]. Type locality: USA, Arizona, 6 miles SE Patagonia. nuda Townsend, 1927: 309 (Eumacrohoughia). Lectotype male (USNM) by fixation of Townsend (1941: 265); this specimen cannot be found in the USNM collection, and is not mentioned by Toma & Nihei (2006) as remaining in São Paulo. Paralectotype female (USNM) [examined by DMW]. Type locality: Brazil, São Paulo, Itaquacetuba. Comb. nov. orbitalis Townsend, 1927: 364 (Verrugomyia). Holotype male (USNM) [examined]. Type locality: Peru, [Lima], Quebrada de Verrugas. Comb. nov. parva Townsend, 1935a: 233 (Pararrhinactia). Holotype male (BMNH) [examined by DMW, including a paratype in USNM]. Type locality: Trinidad. Comb. nov. plagioides van der Wulp, 1890b: 125 (Prospherysa). Lectotype male (BMNH), by designation herein (see Lectotype Designations section). Type locality: Mexico, Morelos, Cuernavaca. Comb. nov. punctiger Townsend, 1927: 346 (Petrargyrops). Lectotype male (USNM) by designation herein (see Lectotype Designations section). Type locality: Brazil, São Paulo, Itaquaquecetuba. Comb. nov. quadra Wiedemann, 1830: 328 (Tachina). Type(s) male (NHMW) [not examined]. Type locality: Brazil. This name was assocated with Pammaerus Aldrich by Townsend (1941: 301) because of the statement “Tachina quadra W may belong here or in Macrohoughia” and, probably following this suggestion, quadra was also included in Pammaerus by Guimarães (1971). Pammaerus is synonymized with Houghia above, and as a result of these actions, T. quadra is therefore tentatively placed in Houghia pending examination of the type(s). Comb. nov. setinervis Coquillett, 1898: 236 (Hypostena). Holotype female (USNM) [examined by DMW]. Type locality: USA, Florida, Biscayne Bay. Combination established by O’Hara & Wood (1998: 762). setipennis Coquillett, 1897: 118 (Houghia). Holotype male (published as female) (USNM) [examined by DMW]. Type locality: USA, Georgia, Tifton. sexualis Curran, 1934b: 469 (Agrarialia). Lectotype male (AMNH), by fixation of Arnaud (1963d: 106) (mention of “Holotype male” from Corozal, Panama in AMNH is regarded as a lectotype fixation) [examined by DMW]. Type locality: Panama, Canal Zone, Corozal. Comb. nov. similis Townsend, 1927: 357 (Sisyrohoughia). Lectotype female (USNM), by fixation of Townsend (1941: 316) (mention of “Ht female” from Itaquaquecetuba in USNM is regarded as a lectotype fixation for the single female syntype in USNM). Type locality: Brazil, São Paulo, Itaquaquecetuba. Comb. nov. simillima Thompson, 1963: 327 (Carceliocephala). Lectotype female (CNC) by designation herein (see Lectotype Designations section). Type locality: Trinidad, Maracas Valley. Comb. nov. sordida van der Wulp, 1890b: 113 (Masicera). Lectotype male (published as female) (BMNH), by designation herein (see Lectotype Designations section). Type locality: Mexico, Tabasco, Teapa. Comb. nov. sternalis Coquillett, 1897: 109 (Sturmia). Holotype male (published as female) (USNM) [examined by DMW]. Type locality: USA, Missouri. Combination established by O’Hara & Wood (1998: 772). persimilis Reinhard, 1943: 90 (Pacidianus). Holotype male (CNC) [examined by DMW]. Type locality: USA, New York, Long Island, Babylon. Combination and synonymy established by Wood in O’Hara & Wood (1998: 772). tenuiseta Macquart, 1846: 292 (164) (Masicera). Lectotype male (published as female) (BMNH) by designation HOUGHIA OF ACG, COSTA RICA Zootaxa 3858 (1) © 2014 Magnolia Press · 11 herein (see Lectotype Designations section). Type locality: Venezuela. Although only the female was described, Macquart’s statement of the length “2 1/2, 3 [lines]” suggested that more than one specimen was involved. The BMNH type catalogue lists LT male and PLT female, Venezuela “Collection de M. Bigot.” Comb. nov. tropica Townsend, 1934: 405 (Tapjohoughia). Holotype male (USNM) [examined by DMW]. Type locality: Brazil, Para, Río Tapajós, Bôa Vista. Comb. nov. Lectotype designations The following lectotypes are designated by DMW in the interests of nomenclatural stability and to restrict each name to a particular specimen. Specimens of the nominal species described by van der Wulp (1890a, 1890b, 1890c) from Mexico each bear 5 labels in addition to a DMW lectotype or paralectotype label, as follows: (1) a yellow-bordered circular white label indicating syntype; (2) a data label, with locality, state, month, and collector; (3) a small label indicating male or female (usually incorrect because van der Wulp was misled by the presence of proclinate orbital setae in males of Houghia); (4) a determination label, prefaced by “B.C.A. [Biologia Centrali-Americana] Dipt. II” and followed by “v. d.W.”; and (5) an accession label “Central America. Pres. by F. D. Godman. O. Salvin. 1903–172”. To avoid repetition, only the data label is given below. The determination label is the same as in van der Wulp’s publications. In his 1927 paper, written in Portuguese, with Portuguese abbreviations, Townsend provided the number of specimens and their localities and dates of collection, It was probably Townsend who labelled one or more of the specimens of the type material with a red card bearing only the word “Type”, and sometimes a sex symbol; otherwise he did not state much other relevant information about the specimens in the 1927 paper. Some of these deficiencies were rectified later in his Manual of Myiology (Townsend 1941) in which he stated the sex and depository of the type specimen of the type species of each genus, which provided at least a guideline in selecting lectotypes. His collection was not sent to USNM until sometime after his death, after the pins and labels of many had seriously deteriorated (N.E. Woodley, pers. comm.). Some remained in Brazil. Red USNM paratype labels with type numbers, which accompany a few but not all the potential syntypes, must have been added after this time by museum staff because Townsend did not return to the United States after he became domiciled in Brazil. Anisia approximata van der Wulp, 1890c: 202. Described from “Two female specimens”; two males are now present in BMNH. Lectotype male (published as female), from Mexico, Guerrero, Venta de Zopilote, 2800', Oct. (H. H. Smith), with a label to be added by DMW: “LECTOTYPE ♂/ of Anisia/ approximata Wulp/ designated 2014/ by D.M. Wood”. Paralectotype one female, same data as lectotype. Paralectotype: one male, same data and depository as lectotype. Actinoprosopa facialis Townsend, 1927: 283. Described from “6 ♂ e 4 ♀, Itaquaquecetuba, S. P., 23 de Ag. até 2 de Out [in the original publication it appears to be Oot or Oct]., em folhagem; in copula, 2 de Set.”; the original syntypes are present in the following institutions: three males and two females in USNM; one male and one female, and one specimen of undetermined sex in MZSP (Toma & Nihei 2006: 251); one male and one female in SENASA. Townsend (1941: 229) mentioned “Ht male” from Itaquaquecetuba in USNM, but this statement cannot be accepted as a lectotype fixation because the specimen in question is not distinguishable from among the three males in USNM. One of the males in USNM is labelled as “Type” and it is hereby designated as lectotype. Lectotype male (USNM), in good condition, labelled: “In Cop ♂/ Itq. [Brazil, São Paulo, Itaquaquecetuba], 2.IX/ On Foliage [in Townsend’s handwriting]// Type [red card]// LECTOTYPE ♂/ of Actinoprosopa/ facialis Tnsd/ designated 2014/ by D.M. Wood”. Paralectotypes 4 males, 4 females, and 1 specimen of undetermined sex, all from same locality as lectotype. In USNM: 1 male, 23.VIII; 1 male, 7.IX; 1 female, 25.VIII, 1 female, 27.VIII; 1 of each sex of the foregoing also labelled “Actinoprosopa/ facialis TT/ [sex symbol] [in Townsend’s handwriting]”. In MZSP (Toma & Nihei 2006: 251): 1 male, 15.IX; 1 female, 25.IX; one specimen of undetermined, sex, 2.X. In SENASA: 1 male and 1 female [the pair of specimens in copula], 2.IX. 12 · Zootaxa 3858 (1) © 2014 Magnolia Press FLEMING ET AL. Masicera impedita van der Wulp, 1890a: 107. Described from “Two female specimens”; one male and one female are now present in BMNH. Lectotype male (published as female) (BMNH), from Mexico, Morelos, Cuernavaca, June (H.H. Smith), with a label to be added by DMW: “LECTOTYPE ♂/ of Masicera/ impedita Wulp/ designated 2014/ by D.M. Wood”. Paralectotype one female, same data as lectotype. Macrohoughia marmorata Townsend, 1927: 325. Described from “Muitos ♂ e ♀, Itaquaquecetuba e Cantareira, S. P., todo o anno, mas especialmente em Ag., em folhagem”. Six of the original syntypes are present in the following institutions: three males and one female in USNM; one male and one female in MZSP (Toma & Nihei, 2006: 251). Lectotype male (USNM), in good condition, labelled: “Itq. [Brazil, São Paulo, Itaquaquecetuba], 27.VIII/ On Foliage// SaoPaulo/ Brazil// CHTTownsend/ coll.// Macrohoughia/marmorata TT/ DetCHTT ♂ [in Townsend’s handwriting]”// ParatypeNo./41690// LECTOTYPE ♂/ of Macrohoughia/ marmorata Tnsd/ designated 2014/ by D.M. Wood”. Paralectotypes, 4 males and 2 females, all from same locality as lectotype. In USNM: 2 males and 1 female; 1 male 1.IX, 1 male 26.VIII, 1 female 18.II., labelled: “Macrohoughia/ [generic name crossed out]/ marmorata TT/ DetCHTT [in Townsend's handwriting]”. In MZSP: 1 male, 25.VIII, and 1 female, 23.VIII. Eumacrohoughia minor Thompson, 1963: 333. Described, apparently from two specimens, from: Male: Trinidad, “Brazil” [village]; II.18.61; Female: Trinidad, Arima Valley; III.9.61.” but without mention of either as holotype. These two syntypes are present in CNC. Cooper & O’Hara (1996) mentioned a male syntype with a red holotype label from Trinidad, Brazil [village], but this holotype label was attached years after publication, and a lectotype was not fixed. Lectotype male (CNC), in poor condition, labelled: “HOLOTYPE [red card]/ Eumacrohoughia/ minor/ Thomp./ CNC No. 9791// Brazil/ II.18.61./ Trinidad. BWI.// CNC Syntype [yellow card]/ Eumacrohoughia/ minor Thompson/ Label affixed 1994// LECTOTYPE ♂/ of Eumacrohoughia/ minor Thompson/ designated 2014/ by D.M. Wood”. Paralectotype, 1 female, labelled: “ALLOTYPE [red card]/ Eumacrohoughia/ minor/ Thomp./ CNC NO. 9791/ / Arima V./ TRINIDAD W.I./ III. 9. 1961 W. R. Thompson// 19.01.III 2911// CNC Syntype [yellow card]/ Eumacrohoughia/ minor Thompson/ Label affixed 1994”. Prospherysa plagioides van der Wulp, 1890a: 105. Described from “Several female specimens” from two localities in Mexico, one from Morelos, Cuernavaca, the remainder from Guerrero, Amula, 6000 feet. Four of the original syntypes are present in BMNH and two in ZMAM. Lectotype male (published as female) (BMNH), in poor condition, from Mexico, Morelos, Cuernavaca, June (H.H. Smith), with a label to be added by DMW: “LECTOTYPE ♂/ of Prospherysa/ plagoiodes Wulp/ designated 2014/ by D.M. Wood”. Paralectotypes, 4 males and 1 female. In BMNH: 2 males, 1 female (abdomen in gelatine capsule), Guerrero, Amula. In ZMAM: 2 males, Guerrero, Amula. According to earlier notes by DMW they may not be conspecific with the lectotype. Petrargyrops punctiger Townsend, 1927: 346. Described from “8 ♂, Itaquaquecetuba, S. P., 10 de Set. até 30 de Nov. em folhagem.” Thompson (1963: 338) treated the species but did not single out a particular specimen as lectotype. Five of the syntypes are present in the following institutions; in USNM, three males, in MZSP, one male, in CNC, one male. In BMNH, one specimen, possibly a syntype. Lectotype male (USNM), in good condition, labelled: “Itq. [Brazil, São Paulo, Itaquaquecetuba], 7.X/ On Foliage// Type [red card]// Townsend/ Genotype/ Collection// Petrargyrops/ punctiger/ DetCHTT ♂ [in Townsend’s handwriting]// LECTOTYPE ♂/ of Petrargyrops/ punctiger Tnsd/ designated 2014/ by D.M. Wood”. Paralectotypes, 4 males, all from same locality as lectotype. In USNM: 1 male 10.IX, 1 male 11.IX, each with additional label added later by museum staff (Townsend’s types were sent to USNM after Townsend’s death) HOUGHIA OF ACG, COSTA RICA Zootaxa 3858 (1) © 2014 Magnolia Press · 13 “ParatypeNo./ 41687/ U.S.N.M. (on red card)”. In MZSP: 1 male, 1.IX (Toma & Nihei 2006: 250). In CNC: 1 male, 30.XI (from Villeneuve/Mesnil collection, with Townsend’s determination label but not labelled as type). Carceliocephala simillima Thompson, 1963: 327. Described from “three females” from “Trinidad, Maracas Valley, X.9.53”. Two females are now present in CNC. Numbers below each specimen apparently refer to a notebook and to slides, the latter of which have deteriorated badly, of eggs and first instar larvae. Lectotype female (CNC), in fair condition, labelled: “Maracas V./ Trinidad, BWI./ Nov-11-1952// Collector/ F. J. Simmonds// 929// 2// Carceliocephala/ simillima nsp// LECTOTYPE ♀/ of Carceliocephala/ simillima Thomps./ designated 2014/ by D.M. Wood”. Paralectotype, 1 female, labelled: “I. C. T. A. [Imperial College of Tropical Agriculture, now University of the West Indies]/ Trinidad, BWI./ Oct. 9–1953// Collector/ F. J. Simmonds// 1224/ II – 2164// PARALECTOTYPE ♀/ of Carceliocephala/ simillima Thomps./ designated 2014/ by D.M. Wood”. We assume that this specimen is an original syntype and that Thompson (1963) overlooked the locality in his original description. Masicera sordida van der Wulp, 1890b: 113. Described from “Several female specimens” from three localities in Mexico. Seven of the original syntypes are now present in BMNH, one from Guerrero, Venta de Zopilote, one from Veracruz, Atoyac, and five from Tabasco, Teapa. The specimen from Venta de Zopilote appears to belong to Lespesia Robineau-Desvoidy. The specimen from Atoyac, although belonging to Houghia, is a female, thus still unidentifiable to species. One of the five specimens from Teapa belongs to Hyphantrophaga Townsend, but the remaining four from Teapa consist of two males each of two different species of Houghia. One of these has been designated as lectotype. Lectotype male (published as female) (BMNH), in fair condition, from Mexico, Tabasco, Teapa (H H. Smith), with a label to be added by DMW: “LECTOTYPE ♂/ of Masicera/ sordida Wulp/ designated 2014/ by D.M. Wood”. Paralectotypes, 4 males, 1 female and 1 without record of sex; 1 male with same data as lectotype and conspecific, 2 males with same data as lectotype belonging to Houghia but not conspecific, 1 male with same data as lectotype belonging to Hyphantrophaga, 1 specimen from Venta de Zopilote belonging to Lespesia, and 1 female from Atoyac belonging to Houghia sp. All, in addition to the locality and accession labels described, are labelled “PARALECTOTYPE/ of Masicera/ sordida Wulp/ designated 2014/ by D.M. Wood”. The lectotype of Masicera sordida was compared to the new species of Houghia described herein. It was thought to be similar to H. omissa, however on further examination it was found to be morphologically different due to the presence of a normal sex patch on tergite 5, a trait not present in H. omissa. Masicera tenuiseta Macquart, 1846: 292 (163). Described from an unstated number of specimens from Venezuela, as female. Macquart’s statement of length “2 1/2, 3 lines” suggests more than one specimen was included. One male and 1 female are now present in BMNH, acquired with the Bigot collection. Lectotype male (published as female) (BMNH), labelled: “Masicera tenuiseta Macq. n.sp. [in Macquart’s handwriting]// Masicera/ tenuiseta Macq.// SYNTYPE ♂, VENEZUELA ex. Bigot Coll. B.M.1960–539 [in Crosskey’s handwriting]// M. tenuiseta? ♀// Amer. Merid. Macq. [in Bigot's handwriting]// LECTOTYPE ♂/ of Masicera/ tenuiseta Macq./ designated 2014/ by D.M. Wood”. Paralectotype, 1 female, same locality as lectotype. Diagnosis of the Genus Houghia The genus Houghia was keyed out in the Manual of Nearctic Diptera under the name Eumasicera, in couplet 27 (Wood 1987: 1206) and in the Manual of Central American Diptera as Houghia in couplets 43 and 49 (Wood & Zumbado 2010: 1362). As in all other Goniini the three postsutural supra-alar setae are well-developed and evenly spaced, with the anteriormost being stouter than the first postsutural dorsocentral seta. The parafacial, katepimeron, hind coxa, and the upper half or more of the facial ridge are bare. Abdominal tergites lack median discal setae. The three major setae of the postpronotum are arranged in a triangle. The ocellar setae are always proclinate and in most 14 · Zootaxa 3858 (1) © 2014 Magnolia Press FLEMING ET AL. species arise beside the anterior ocellus (Fig. 5). Males of all species have two pairs of well-developed proclinate fronto-orbital setae as in females, and this character state distinguishes them from males of Hyphantrophaga and Carcelia in which they are absent. Except for the four species in couplets 27 and 28 below, in which the height of the gena is about one-tenth the height of the head, which approaches the state in some Hyphantrophaga, the height of the gena is one-twelfth or less the height of the head, thus resembling members of Carcelia. The eye of a few species is conspicuously haired (Fig. 6b) but in most species these hairs are sparse and inconspicuous or apparently absent. The external morphological traits used to separate species are given in the following list. Table 2 presents a comprehensive list of external and internal characters and their relative states in each species. The characters are presented in order of appearance on the body anteriorly to posteriorly and arranged by head, thorax, and abdomen. Head. 1a. Antenna black. 1b. Pedicel and usually upper part of first flagellomere pale orange (Fig. 1). FIGURE 1. Frontal view of head, detailing the difference in colour at the base of the antennae. a. Houghia sexmaculata sp. nov. b. Houghia fimbriata sp. nov. 2a. Base of antenna, when viewed in profile, distinctly above middle of eye, distance between vibrissal and antennal angles usually greater than distance between antennal angle and vertex, or eye elongate (Fig. 2a). 2b. Base of antenna inserted approximately at mid-eye level; eye more rounded (Fig. 2b). 3a. Length of first flagellomere extending to facial margin. 3b. Length of first flagellomere almost extending to facial margin (usually shorter by less than the length of the pedicel). 3c. Length of first flagellomere shorter than facial margin (usually shorter by at least the length of the pedicel). 4a. Facial ridge bare except for a few (usually 3–5) decumbent small setae above vibrissa. 4b. Facial ridge with decumbent setae and a row of minute, inconspicuous, hairs (easily overlooked) on lower half or more (Fig. 3). 5a. Palpus pale, usually distinctly yellowish. 5b. Palpus dark, gray to black. 6a. Postgena, behind postoccipital row, above level of lower facial margin without black setae. 6b. Postgena, behind postoccipital row, above level of lower facial margin with a small patch of black setae. 6c. Postgena, behind postoccipital row, above level of lower facial margin extensively covered in black setae. HOUGHIA OF ACG, COSTA RICA Zootaxa 3858 (1) © 2014 Magnolia Press · 15 16 · Zootaxa 3858 (1) © 2014 Magnolia Press FLEMING ET AL. HOUGHIA OF ACG, COSTA RICA Zootaxa 3858 (1) © 2014 Magnolia Press · 17 18 · Zootaxa 3858 (1) © 2014 Magnolia Press FLEMING ET AL. HOUGHIA OF ACG, COSTA RICA Zootaxa 3858 (1) © 2014 Magnolia Press · 19 20 · Zootaxa 3858 (1) © 2014 Magnolia Press FLEMING ET AL. HOUGHIA OF ACG, COSTA RICA Zootaxa 3858 (1) © 2014 Magnolia Press · 21 22 · Zootaxa 3858 (1) © 2014 Magnolia Press FLEMING ET AL. HOUGHIA OF ACG, COSTA RICA Zootaxa 3858 (1) © 2014 Magnolia Press · 23 24 · Zootaxa 3858 (1) © 2014 Magnolia Press FLEMING ET AL. FIGURE 2. Lateral view of head, detailing the position of the antennae relative to the middle of the eye. a. Houghia romeroae sp. nov. b. Houghia chavarriae sp. nov. 7a. Parafacial silver (Fig. 4a). 7b. Parafacial pale brassy to gold (Fig. 4b). 8a. Fronto-orbital plate pale brassy to gold, on its entire length from vertex to base of antenna. 8b. Fronto-orbital plate gold on upper half, from vertex to lower proclinate orbital seta. 8c. Fronto-orbital plate gold only at vertex, adjacent to ocellar triangle, and the remainder silver. 8d. Fronto-orbital plate with no gold, entirely silver. 9a. Ocellar triangle appearing to be notched anteriorly when viewed from above; gold tomentosity of apex, visible laterally, changes to brown medially, matching frontal vitta, in dorsal view (Fig. 5a). 9b. Ocellar triangle rounded anteriorly (Fig. 5c). 9c. Ocellar triangle sharply pointed anteriorly (Fig. 5b). 9d. Ocellar triangle when viewed from above appearing to be concolorous with frontal vitta. 10a. Ocellar setae arising beside anterior ocellus (Fig. 5c). 10b. Ocellar setae arising behind anterior ocellus but closer to anterior ocellus than to posterior ocelli (Fig. 5b). 10c. Ocellar setae arising distinctly behind anterior ocellus, approximately midway between anterior ocellus and posterior ocelli (Fig. 5a). 11a. Diameter of anterior ocellus approximately equal to diameter of base of adjacent ocellar seta. 11b. Anterior ocellus reduced in size, its diameter less than diameter of base of adjacent ocellar seta. 12a. Eye apparently bare, often a few sparse, inconspicuous, short hairs, shorter than diameter of two ommatidia, usually present but discernment requiring careful observation under high magnification (Fig. 6a). HOUGHIA OF ACG, COSTA RICA Zootaxa 3858 (1) © 2014 Magnolia Press · 25 12b. Eye with more conspicuous hairs, denser and each longer than diameter of two ommatidia (Fig. 6b). 13a. Fronto-orbital plate with small recumbent hairs over most of its surface, usually forming an irregular row next to eye margin between anterior and posterior proclinate orbital setae. 13b. Fronto-orbital plate almost bare, with few hairs, none concentrated adjacent to eye margin. FIGURE 3. Lateral view of head of Houghia fimbriata sp. nov. Arrow indicates the small decumbent hairs along the facial ridge. FIGURE 4. Frontal view comparing gold coverage on fronto-orbital plate. a. Houghia sexmaculata sp. nov. b. Houghia aurifera sp. nov. 26 · Zootaxa 3858 (1) © 2014 Magnolia Press FLEMING ET AL. FIGURE 5. Dorsal view of head. Red lines indicate the shape of the ocellar triangle as described, and placement of anterior ocellus (green circle) relative to the base(s) of ocellar setae (blue circles). a. Houghia marini sp. nov. b. Houghia graciloides sp. nov. c. Houghia griseifrons sp. nov. FIGURE 6. Dorsal view of head detailing eye hairs. a. Eye bare, Houghia fimbriata sp. nov. b. Eye haired, Houghia brevipilosa sp. nov. Thorax. 14a. Postpronotal setae 5, with two small anterior setae in addition to the three main setae. 14b. Postpronotal setae restricted to the three main ones. 15a. Dark stripes on scutum on either side of dorsocentral row of setae separated by yellow tomentosity concolorous with that of remainder of scutum. 15b. Tomentosity between median and lateral stripes brownish or gray, paler than dark stripes, but darker than remainder of scutum. 16a. Median and lateral stripes on either side of scutum separate from each other. 16b. These stripes united to one another posteriorly. HOUGHIA OF ACG, COSTA RICA Zootaxa 3858 (1) © 2014 Magnolia Press · 27 17a. Scutum with 4 pairs of postsutural dorsocentral setae (Fig. 7a). 17b. Scutum with only 3 pairs of postsutural dorsocentral setae, with a gap between first and third seta (Fig. 7b). 18a. Anterior quadrant of anepisternum covered with short setae except for 3 to 5 distinctly larger setae, usually forming an oblique row. 18b. This area covered with long setae, none distinctly larger than the others or forming a row. FIGURE 7. Dorsal view of thorax showing differing number of postsutural dorsocentral setae. Red circles indicate positions of these setae. a. Houghia fimbriata sp. nov., with four postsutural dorsocentral setae. b. Houghia omissa sp. nov., with three postsutural dorsocentral setae. FIGURE 8. Lateral view of thorax. Red circle indicates position of middle katepisternal seta. a. Houghia pilosifrons sp. nov., with middle katepisternal seta present but small. b. Houghia sexmaculata sp. nov., middle katepisternal seta absent. 28 · Zootaxa 3858 (1) © 2014 Magnolia Press FLEMING ET AL. 19a. Katepisternum with three setae, the middle one always the smallest (Fig. 8a). 19b. Katepisternum with only two setae (Fig. 8b). 20a. Vein R1 bare dorsally (Fig. 9a). 20b. Vein R1 with a row of small recumbent setae along its entire length (Fig. 9b). 21a. Legs ranging from reddish brown to yellow tinged but overall dark. 21b. Legs distinctly pale, often yellow. 22a. Coxae dark usually concolorous with remainder of legs. 22b. Coxae distinctly yellow tinged, contrasting with darker remainder of legs. FIGURE 9. Dorsal view of wing, showing haired nature of vein R1. a. Bare, Houghia bivittata sp. nov., position of vein R1 shown in red. b. Haired, Houghia nigrofemur sp. nov. Abdomen. 23a. Ground colour of dorsal surface of abdomen black. 23b. Ground colour of dorsal surface of tergite 3 and in some species parts of other tergites as well, dark medially but reddish laterally. 24a. Ground colour of ventral surface of abdomen entirely black. 24b. Ground colour of ventral surface partially or entirely reddish or yellowish. 25a. Sex patches present on tergite 4 and usually on tergite 5 (although much smaller) (Figs. 10b,d,e). 25b. Sex patches present on tergite 3, as well as on tergites 4 and 5 (though sometimes small or absent on tergite 5) (Fig. 10a). HOUGHIA OF ACG, COSTA RICA Zootaxa 3858 (1) © 2014 Magnolia Press · 29 25c. Sex patches present only on tergite 4 (Fig. 10c). 25d. Sex patches absent. 26a. Ground colour beneath sex patches shiny black. 26b. Ground colour beneath sex patches not distinctly shiny. FIGURE 10. Ventral view of abdomen showing different configurations of sex patches. a. Houghia graciloides sp. nov., showing characteristic sex patch on tergites 3, 4, and 5. b. Houghia longipilosa sp. nov., with sex patch present on tergites 4 and 5 only. c. Houghia omissa sp. nov., with sex patch present on tergite 4 only. d. Houghia confinis sp. nov with sex patch on tergites 4 and 5. e. Houghia macilenta sp. nov., with no sex patch at all. Key to Males of Houghia Reared from Caterpillars in ACG, Northwestern Costa Rica 1 2 3 4 - Sex patches (areas of shorter, denser, appressed hairs, often referred to as sex patches or Sturmia spots, present only in males) lacking, undersides of tergites 4 and 5 tomentose as on dorsal surfaces; vein R1 setose along its entire length dorsally. . . . . . . 2 Sex patches present (Fig. 10); if in doubt undersurface of tergite 4 differing in appearance from dorsal surface; vein R 1 bare dorsally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Scape and upper third or more of first flagellomere pale orange; first flagellomere slightly to strongly curved or sinuous (Figs. 35c, 36c), its length surpassing lower facial margin by more than length of pedicel; dark stripes on scutum distinct (Figs. 35a, 36a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Antenna entirely black except for slightly paler base of first flagellomere; first flagellomere straight, its length barely surpassing lower facial margin; dark scutal stripes indistinct (Figs. 25a, 32a). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Bases of femora orange, contrasting with dark apices of femora, tibiae and tarsi; first flagellomere distinctly S-shaped, the lower curve parallelling strongly curved facial ridge (Fig. 36c). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ochrofemur sp. nov. Legs entirely black; first flagellomere merely slightly bowed anteriorly (only one male available) . . . . . . nigrofemur sp. nov. Lateral margins of frontal vitta yellowish tomentose, leaving only a narrow dark median line (Fig. 32b) and appearing to connect ocellar triangle to pale golden fronto-orbital plates. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . macilenta sp. nov. Frontal vitta wider, entirely brown, separating ocellar triangle from grayish fronto-orbital plates . . . . . . . . griseifrons sp. nov. 30 · Zootaxa 3858 (1) © 2014 Magnolia Press FLEMING ET AL. 5 6 7 8 9 10 11 12 13 14 - 15 16 - 17 18 19 20 21 22 - Sex patches present on black integument on tergites 3 as well as on tergites 4 and 5; palpus dark gray to black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .sexmaculata sp. nov. Sex patches present on tergite 4, and usually also on tergite 5, but if apparently also on tergite 3 (as in H. gracilis (Fig. 10a) and H. graciloides) then integument on ventral surface reddish; palpus pale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Pedicel and upper part of first flagellomere pale orange . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 Pedicel and first flagellomere entirely dark . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 Facial ridge with row of fine white hairs on at least lower half, easily overlooked and best viewed in profile against a dark background; postgena posterior to postoccipital fringe, with a small patch of black setae . . . . . . . . . . . . . . . fimbriata sp. nov. Facial ridge bare except for the usual few supravibrissal setae; postgena posterior to postoccipital fringe lacking black setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . pallida sp. nov. Eye conspicuously haired, each hair at least as long as width of three or more ommatidia together; setae of anterodorsal quadrant of anepisternum increasing in length anteriorly but lacking row of 3 or 4 distinctly larger setae. . . . . . . . . . . . . . . . . . . . . 9 Eye bare or apparently so except at higher magnification, hairs if present, shorter than width of two ommatidia; setae of anterodorsal quadrant of anepisternum short except for row of 3 or 4 distinctly longer and thicker setae . . . . . . . . . . . . . . . . . . . . . 11 Parafacial and fronto-orbital plate entirely golden tomentose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . aurifera sp. nov. Parafacial silvery, contrasting with gold of vertex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 Posterior dark bands of abdominal tergites 2–5 grayish-brown tomentose, indistinctly differentiated from paler tomentum of remaining anterior portion of tergites; fronto-orbital plate gold pollinose only to level of anterior proclinate fronto-orbital seta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . brevipilosa sp. nov. Posterior bands of abdominal tergites black, clearly differentiated from paler anterior portions of tergites; fronto-orbital plate gold tomentose to base of antenna . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . longipilosa sp. nov. Scutum with 3 pairs of postsutural dorsocentral setae, with a gap between the first seta and the next (suggesting a missing seta) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 Scutum with 4 pairs of postsutural dorsocentral setae, more or less equally spaced . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 Tergite 5 yellowish tomentose ventrally, lacking any trace of sex patch, contrasting with well developed patch on tergite 4 (Fig. 10c) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . omissa sp. nov. Tergite 5 with sex patch, similar to, although smaller than, patch on tergite 4, on black integument . . . . . . . . . . . . . . . . . . . . 13 Scutum with longitudinal dark stripes on either side of dorsocentral row of setae connected to one another by dark brown tomentosity, creating the appearance of a single wide stripe on each side; integument of abdominal tergite 3 more reddish laterally than adjacent tergites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 Scutum with 4 (2 pairs of) narrow longitudinal dark stripes separated from each other by golden tomentosity concolorous with rest of scutum; abdominal tergite 3 concolorous with adjacent tergites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 Fronto-orbital plate gold tomentose from vertex to antennal bases; ocellar setae arising behind anterior ocellus; colour of brown tomentosity surrounding dorsocentral setae nearly as dark as longitudinal stripes . . . . . . . . . . . . . . . . . bivittata sp. nov. Fronto-orbital plate silvery tomentose except for a tinge of gold extending from vertex to anterior proclinate fronto-orbital seta; ocellar setae arising slightly in front of anterior ocellus; colour of brown tomentosity surrounding dorsocentral setae paler than longitudinal stripes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . biseriata sp. nov. Inner and outer dark longitudinal stripes on each side of scutum converging posteriorly to coalesce at level of second last postsutural dorsocentral seta, thus enclosing, or nearly enclosing, seta with darker tomentosity . . . . . . . . . . . . . . . confinis sp. nov. Inner and outer dark stripes remaining separate posteriorly separated by yellow tomentum concolorous with rest of scutum. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 Ocellar triangle, observed from above, sharply pointed anteriorly, the point extending to level of upper fronto-orbital seta; gold tomentosity of fronto-orbital plate extending forward only to same level; first flagellomere extending to lower facial margin. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . triangularis sp. nov. Ocellar triangle rounded, truncate, or notched anteriorly (point concolorous with vitta hence scarcely distinguishable), extending forward only to level of upper reclinate frontal seta; gold tomentosity of fronto-orbital plate extending forward to or nearly to level of upper fronto-orbital seta; first flagellomere shorter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 Height of apical portion (fused part to apex) of cerci three times height of dorsal notch, enclosing anus (Fig. 19d); surstylus dagger-shaped, pointed apically (Fig. 19e) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . crypta Townsend Height of fused portion of cerci less than twice height of dorsal notch (Fig, 21d); surstylus bluntly rounded apically . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . destituta sp. nov. Ground colour of ventral surface of abdominal tergites 1 to 4 yellow or orange, somewhat translucent. . . . . . . . . . . . . . . . . . 19 Ground colour of ventral surface of abdomen entirely black, usually obscured by paler tomentosity. . . . . . . . . . . . . . . . . . . . 23 Katepisternal setae 2 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . luteiventris sp. nov. Katepisternal setae 3 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 Sex patches on tergites 3, 4, and 5 consisting of shorter more appressed hairs than elsewhere on abdomen, but these hairs sparse and indistinct because of reddish background; ocellar setae arising behind anterior ocellus . . . . . . . . . . . . . . . . . . . . . 21 Sex patches confined to tergites 4 and 5; ocellar setae arising beside anterior ocellus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 Eye apparently bare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . gracilis sp. nov. Eye with minute obscure hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . graciloides sp. nov. Fronto-orbital plate mostly silvery, with pale golden tomentosity, and area of small hairs, extending from vertex only to level of anterior fronto-orbital seta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . delospilota sp. nov. Fronto-orbital plate entirely golden tomentose and more extensively haired, extending from vertex to lower margin of pedicel . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . pilosifrons sp. nov. (in part) HOUGHIA OF ACG, COSTA RICA Zootaxa 3858 (1) © 2014 Magnolia Press · 31 23 24 25 26 27 28 29 30 31 32 33 34 35 - Fronto-orbital plate entirely gold tomentose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24 Gold tomentosity of fronto-orbital plate restricted to vertex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 Parafacial silver tomentose, changing abruptly at level of antenna to gold . . . . . . . . . . . . . . . . . . . . . . . . . . pilosifrons (in part) Parafacial pale yellow tomentose gradually changing at level of antenna to darker brassy yellow on fronto-orbital plate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . aerata sp. nov. Height of gena one-tenth height of head in lateral view . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 Height of gena one-twelfth or less height of head . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29 Depth of V-shaped dorsal notch of cerci in posterior view one quarter or less height of entire cerci (Figs. 30d, 28d); lateral scutal stripe converging to meet medial stripe at its posterior end (Figs. 30a, 28a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27 Depth of V-shaped notch one third or more height of cerci (Figs. 27d, 45d); both pairs of scutal stripes separated along their entire lengths (Figs. 27a, 45a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28 Dorsal lobe of syncerus in posterior view almost as broad as high, bluntly rounded dorsally, depth of V-shaped dorsal notch one quarter height of cerci (Fig. 28d); apices of cerci splayed apart apically. . . . . . . . . . . . . . . . . . . . . . . . . . . latilobus sp. nov. Dorsal lobe of cerci more pointed and attenuated dorsally; depth of V-shaped notch less than one quarter (about 1/4.5) height of cerci (Fig. 30d); apices of cerci straight and parallel . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . longicercus sp. nov. Cerci in posterior view slightly constricted at midlength (Fig. 27d); posteroventral quadrant of anepimeron sparsely short haired. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . latigena sp. nov. Cerci in posterior view tapering evenly to apex (Fig. 45d); posteroventral quadrant of anepimeron more densely long haired . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . velutina sp. nov. Width of frontal vitta one-tenth width of broad shiny silvery fronto-orbital plate . . . . . . . . . . . . . . . . . . . . . . . parmata sp. nov. Width of frontal vitta greater than one-tenth of fronto-orbital plate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30 Gold tomentosity of fronto-orbital plate confined to vertex, not extending forward farther than upper fronto-orbital setae. . . 31 Gold tomentosity extending at least to lower fronto-orbital setae and often to the antennal bases . . . . . . . . . . . . . . . . . . . . . . 34 First flagellomere extending to lower facial margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32 First flagellomere terminating above lower facial margin by about half length of pedicel . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33 Surstylus distinctly longer than cerci (Figs. 26 d, e), its apex surpassing that of apex of cerci by more than the width of the surstylus; [palpus of female swollen apically] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . inflatipalpis sp. nov. Surstylus barely longer than cerci (Figs. 33, d, e); [palpus of female as in all other species (except H. luteipennis), only slightly enlarged apically] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . marini sp. nov. Gold tomentosity of fronto-orbital plate restricted to upper margin, not extending forward beyond upper reclinate orbital seta and apex of ocellar triangle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . blancoi sp. nov. Gold tomentosity more extensive, extending to upper fronto-orbital seta . . . . . . . . . . . . . . . . . . . . . . . . . . . . romeroae sp. nov. Surstylus long, straight, and parallel-sided in lateral view (Fig. 43e); apical processes of sternite 5 elongate, slender, and directed medially (Fig. 43f). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . spathulata sp. nov. Surstylus in lateral view obovate, widened moderately at mid-length; apical processes of sternite 5 shorter, thicker and directed posteriorly (Figs. 17f, and 34f) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35 Posterior margin of surstylus rounded in lateral view (Fig. 17d) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . chavarriae sp. nov. Posterior margin of surstylus slightly concave in lateral view (Fig. 34d). . . . . . . . . . . . . . . . . . . . . . . . . . . . .matarritai sp. nov. Species diagnoses and descriptions Houghia aerata Fleming & Wood, sp. nov. Fig. 11 a–f Diagnosis. One of two species (the other being H. aurifera) with gold tomentose head, darker on the frons, slightly paler on the parafacial. In H. aerata, however, the eye hairs are short and inconspicuous, unlike the longer eye hairs of aurifera. Description. Male. Antenna black. When viewed in profile, antenna arises approximately at level of middle of eye. Length of first flagellomere almost extending to facial margin (usually shorter by less than length of pedicel). Facial ridge bare except for a few (usually 3–5) decumbent small setae above vibrissa. Palpus pale, usually distinctly yellowish. Postgena behind postoccipital row, above level of lower facial margin, with a small patch of few black setae. Parafacial pale brassy to gold. Colour of fronto-orbital plate pale brassy to gold on its entire length from vertex to base of antenna (more than 50% coverage). Surface of fronto-orbital plate covered with small recumbent hairs, especially near margin of eye. Ocellar triangle, when viewed from above appearing to be notched anteriorly. Diameter of anterior ocellus equal to, or greater than, diameter of base of adjacent ocellar seta. Ocellar setae arising beside, or slightly in front of, anterior ocellus. Eye bare. Postpronotum with 4 or 5 postpronotal setae. Dark stripes on either side of dorsocentral row of setae, separated from one another by yellow tomentosity. Median and lateral stripes on either side of scutum separate from each other posteriorly. Postsutural dorsocentral setae 4. 32 · Zootaxa 3858 (1) © 2014 Magnolia Press FLEMING ET AL. Anterior quadrant of anepisternum covered with short setae except for usually 3 to 5 distinctly larger setae. Katepisternum bearing three setae, the middle one always the smallest. Vein R1 bare dorsally. Legs ranging from reddish brown to yellow tinged but overall dark. Coxae dark usually concolourous with remainder of leg. Ground colour of dorsal surface of abdomen dark to black. Ground colour of ventral surface of abdomen entirely black. Sex patches present on tergites 4 and 5. Ground colour of sex patches shiny black. Terminalia: surstylus equilaterally oblong shaped, posterodorsal half bare, apex bearing few short apical spines, tip with strong inwardly apical curve when viewed dorsally. Cerci sharply pointed or distinctly tapered, apex with straight tip, ventral surface haired, separation between cerci straight, up to 85% as long as surstylus. Lobe of sternite 5 small and pointed apically, inner margin covered in dense tomentosity appearing darker than surrounding cuticle, internal edge inwardly curved, apical seta absent. FIGURE 11. General morphology of Houghia aerata sp. nov. a. Dorsal habitus. b. Frontal view of head. c. Lateral habitus. d. Posterior view of male terminalia. e. Lateral view of male terminalia. f. Ventral view of sternite 5 + segments 7+8. Hosts. Houghia aerata has been reared only from a sample of 300+ rain forest Macrocneme cabimensis Dyar (11X) and Macrocneme iole Druce (4X) (Arctiinae, Erebidae) feeding on foliage of vines in the Apocynaceae. Holotype. ♂, CNC. Type locality: Costa Rica, Area de Conservación Guanacaste, Prov. Guanacaste, Sector Pitilla, Pasmompa (11.01926°, -85.40997°), 440 m, 06/09/2004, Petrona Rios, DHJPAR0011496. Paratypes. 6 ♂, 7 ♀ (CNC) Costa Rica, Prov. Alajuela and Guanacaste, ACG database codes: DHJPAR0029683, DHJPAR0008537, DHJPAR0027905, DHJPAR0035906, DHJPAR0011500, DHJPAR0008536, DHJPAR0035889, DHJPAR0035859, DHJPAR0035860, DHJPAR0011483, DHJPAR0008538, DHJPAR0016119. HOUGHIA OF ACG, COSTA RICA Zootaxa 3858 (1) © 2014 Magnolia Press · 33 Etymology. From the Latin adjective, aeratus, meaning “of copper, bronze, or brass”, in reference to the brassy colour of the tomentosity covering all parts of the head, especially the fronto-orbital plate and parafacial. Distribution. Costa Rica, ACG, Prov. Alajuela & Guanacaste, rain forest, 320–700 m elevation. Houghia aurifera Fleming & Wood, sp. nov. Fig. 12 a–f Diagnosis. The presence of long eye hairs and bright gold tomentosity on the entire head, darker and more golden on the fronto-orbital plate, distinguish this species. Houghia aerata also has a gold head, but paler and more yellowish on face and lower part of parafacial and the eye hairs are minute. Houghia brevipilosa and H. longipilosa, also with long haired eyes, have the head mostly silver, with gold tomentosity confined to the frons. Description. Male. Antenna black. When viewed in profile, antenna arises distinctly above middle of eye. Length of first flagellomere shorter than facial margin, (usually shorter by at least the length of the pedicel). Facial ridge bare except for a few (usually 3–5) decumbent small setae above vibrissa. Palpus pale, usually distinctly yellowish. Postgena behind postoccipital row, above level of lower facial margin, extensively covered with setae, and fine hairs. Parafacial pale brassy to gold. Colour of fronto-orbital plate pale brassy to gold on its entire length from vertex to base of antenna (more than 50% coverage). Surface of fronto-orbital plate covered with small recumbent hairs, especially near margin of eye. Ocellar triangle, when viewed from above appearing rounded anteriorly. Diameter of anterior ocellus equal to, or greater than, diameter of base of adjacent ocellar seta. Ocellar setae arising beside, or slightly in front of, anterior ocellus. Eye with conspicuous hairs. Postpronotum with 4 or 5 postpronotal setae. Dark stripes on either side of dorsocentral row of setae separated from one another by yellow tomentosity. Median and lateral stripes on either side of scutum separate from each other posteriorly. Postsutural dorsocentral setae 4. Anterior quadrant of anepisternum covered with long setae, none distinctly larger than the others or forming a row. Katepisternum with three setae, the middle one always the smallest. Vein R1 bare dorsally. Legs ranging from reddish brown to yellow tinged but overall dark. Coxae dark usually concolourous with remainder of leg. Ground colour of dorsal surface of abdomen dark to black. Ground colour of ventral surface of abdomen entirely black. Sex patches present on tergites 4 and 5. Ground colour of sex patches shiny black). Terminalia: surstylus bayonet shaped, posterodorsal half bare, apex bearing many stout apical spines, tip with light inwardly apical curve when viewed dorsally. Cerci rounded, apex with blunt, hooked tip, ventral surface haired, separation between cerci straight, up to 85% as long as surstylus. lobe of sternite 5 small and pointed apically, inner margin covered in dense tomentosity appearing darker than surrounding cuticle, internal edge inwardly curved, single long apical seta emanating from apex. Hosts. Houghia aurifera has been reared once from each of 14 species of caterpillars in 9 families, and 45 times from small hesperiine and eudamine Hesperiidae feeding on herbaceous plants and small woody plants, in ACG rain forest and dry forest, out of a sample of 15000+ caterpillars. While this appears to be a “generalist”, its somewhat variable DNA barcode suggests the need for further exploration for the presence of cryptic species. Holotype: ♂, CNC. Type locality: Costa Rica, Area de Conservación Guanacaste, Prov. Guanacaste, Sector Mundo Nuevo, Vado Miramonte (10.77175°, -85.43400°), 305 m, 04/15/2007, Jose Cortez, DHJPAR0019684. Paratypes. 28 ♂, 37 ♀ (CNC) Costa Rica, Prov. Guanacaste, ACG database codes: DHJPAR0019687, DHJPAR0048633, DHJPAR0034504, DHJPAR0008299, DHJPAR0039257, DHJPAR0006706, DHJPAR0046408, DHJPAR0008308, DHJPAR0010001, DHJPAR0008546, DHJPAR0008221, DHJPAR0008302, DHJPAR0024607, DHJPAR0007097, DHJPAR0008303, DHJPAR0015027, DHJPAR0024572, DHJPAR0024590, DHJPAR0024591, DHJPAR0024589, DHJPAR0024579, DHJPAR0027802, DHJPAR0048408, DHJPAR0048422, DHJPAR0048423, DHJPAR0048412, DHJPAR0048411, DHJPAR0045668, DHJPAR0048555, DHJPAR0048631, DHJPAR0048424, DHJPAR0030230, DHJPAR0045667, DHJPAR0011504, DHJPAR0011497, DHJPAR0008548, DHJPAR0023615, DHJPAR0048427, DHJPAR0049641, DHJPAR0049566, DHJPAR0048376, DHJPAR0008540, DHJPAR0024585, DHJPAR0035716, DHJPAR0019682, DHJPAR0024592, DHJPAR0008304, DHJPAR0008539, DHJPAR0011499, DHJPAR0040720, DHJPAR0049637, DHJPAR0021965, DHJPAR0024587, DHJPAR0019686, DHJPAR0048681, DHJPAR0046442, DHJPAR0048426, DHJPAR0048419, DHJPAR0029865, DHJPAR0049639, DHJPAR0048425, 09-SRNP-12023, 12-SRNP-40498, 08-SRNP-55893. 34 · Zootaxa 3858 (1) © 2014 Magnolia Press FLEMING ET AL. ARIZONA: 1 ♂ “USA AZ Huachuca Mtns./ Ramsey Cyn. Hamburg/ Trail 10.viii.2013/ 31º26.2'N 110º19.2'W/ ca. 6200' J. E. O’Hara// Legs off for DNA/ OH10-08-13-002”. Etymology. From the Latin adjective, aurifera, meaning “gold-bearing”, in reference to the bright golden tomentosity on the fronto-orbital plate and parafacial. Distribution. Costa Rica, ACG, Prov. Guanacaste and Prov. Alajuela, 95–600 m.; USA, southern Arizona. FIGURE 12. General morphology of Houghia aurifera sp. nov. a. Dorsal habitus. b. Frontal view of head. c. Lateral habitus. d. Posterior view of male terminalia. e. Lateral view of male terminalia. f. Ventral view of sternite 5 + segments 7+8. Houghia biseriata Fleming & Wood, sp. nov. Fig. 13 a–f Diagnosis. One of a group of seven closely related species in ACG, and many more in Central and South America, the H. crypta group, in which the second postsutural dorsocentral seta is missing, leaving a gap between first and third setae (H. omissa, H. confinis, H. triangularis, H. destituta, H. crypta, H. biseriata, H. bivittata). Houghia biseriata shares with H. bivittata the presence of a pair of wide dark scutal stripes (Figs. 13a, 14a) as a result of dark brown tomentosity between the two longitudinal dark scutal stripes on each side. The gold tomentosity on the fronto-orbital plate is confined to the vertex, not extending forward beyond anterior proclinate fronto-orbital seta (Fig. 13b), thus distinguishing this fly from H. bivittata. HOUGHIA OF ACG, COSTA RICA Zootaxa 3858 (1) © 2014 Magnolia Press · 35 FIGURE 13. General morphology of Houghia biseriata sp. nov. a. Dorsal habitus. b. Frontal view of head. c. Lateral habitus. d. Posterior view of male terminalia. e. Lateral view of male terminalia. f. Ventral view of sternite 5 + segments 7+8. Description. Male. Antenna black. When viewed in profile, antenna arises, approximately at level of middle of eye. Length of first flagellomere extending to facial margin. Facial ridge bare except for a few (usually 3–5) decumbent small setae above vibrissa. Palpus pale, usually distinctly yellowish. Postgena behind postoccipital row, above level of lower facial margin, with a small patch of few black setae. Parafacial silver. Colour of fronto-orbital plate gold on upper half, from vertex to midway down face (up to 50% coverage). Surface of fronto-orbital plate almost bare. Ocellar triangle, when viewed from above appearing to be notched anteriorly. Diameter of anterior ocellus less than diameter of base of adjacent ocellar seta. Ocellar setae arising beside, or slightly in front of, anterior ocellus. Eye bare. Postpronotum restricted to the three main postpronotal setae. Dark stripes on either side of dorsocentral row of setae, separated from one another by yellow tomentosity. Median and lateral stripes on either side of scutum separate from each other posteriorly. Postsutural dorsocentral setae 3, with a gap between first and third seta. Anterior quadrant of anepisternum covered with short setae except for usually 3 to 5 distinctly larger setae. Katepisternum bearing three setae, the middle one always the smallest. Vein R1 bare dorsally. Legs ranging from reddish brown to yellow tinged but overall dark. Coxae dark usually concolourous with remainder of leg. Ground colour of dorsal surface of abdominal tergite 3 and in some specimens parts of other tergites as well, dark medially but reddish laterally. Ground colour of ventral surface of abdomen partially or entirely reddish or yellowish. Sex patches present on tergites 4 and 5. Ground colour of sex patches shiny black. Terminalia: surstylus equilaterally oblong shaped, posterodorsal half haired, apex bearing few short apical spines, tip with light inwardly 36 · Zootaxa 3858 (1) © 2014 Magnolia Press FLEMING ET AL. apical curve when viewed dorsally. Cerci sharply pointed or distinctly tapered, apex with blunt, hooked tip, ventral surface bare, separation between cerci deep narrow V shape, about as long as surstylus. Lobe of sternite 5 large and rounded apically, inner margin covered in dense tomentosity appearing darker than surrounding cuticle, internal edge inwardly curved, single long apical seta emanating from apex. Hosts. Houghia biseriata has been reared 31 times, from a sample of 1244 caterpillars of Dubiella belpa Evans (20X) and Carystina aurifer (Godman & Salvin) (11X) (Hesperiinae, Hesperiidae) feeding on the foliage of 6 species of rain forest understory small palms (Arecaceae). Holotype. ♂, CNC. Type locality: Costa Rica, Area de Conservación Guanacaste, Prov. Guanacaste, Sector Pitilla, Sendero Naciente (10.98705°, -85.42816°), 700 m, 04/30/2008, Manuel Rios, DHJPAR0024455. Paratypes. 26 ♂, 18 ♀ (CNC) Costa Rica, Prov. Alajuela and Guanacaste, ACG database codes: DHJPAR0045599, DHJPAR0011533, DHJPAR0008785, DHJPAR0021026, DHJPAR0008783, DHJPAR0042554, DHJPAR0010325, DHJPAR0008555, DHJPAR0008554, DHJPAR0019540, DHJPAR0008835, DHJPAR0030162, DHJPAR0029650, DHJPAR0008788, DHJPAR0035779, DHJPAR0034611, DHJPAR0040697, DHJPAR0008513, DHJPAR0016112, DHJPAR0011536, DHJPAR0023652, DHJPAR0008512, DHJPAR0035715, DHJPAR0035831, DHJPAR0035790, DHJPAR0011515, DHJPAR0029698, DHJPAR0011491, DHJPAR0008516, DHJPAR0008517, DHJPAR0008547, DHJPAR0011493, DHJPAR0011494, DHJPAR0011512, DHJPAR0011492, 07-SRNP-32205, 09-SRNP-67518, 05-SRNP-42322 06-SRNP-43586, 08-SRNP-31164. Etymology. From the Latin adverb,”bis”, “bi-” meaning two, and the adjective “seriatus” derived from “series”, meaning row, in reference to the pair of broad dark stripes on the scutum. Distribution. Costa Rica, ACG, Prov. Alajuela & Guanacaste, rain forest and dry forest, 96–900 m elevation. Houghia bivittata Fleming & Wood, sp. nov. Figs. 9 a, 10 e, 14 a–f Diagnosis. One of a group of seven closely related species in ACG, and many more in Central and South America, the H. crypta group, in which the second postsutural dorsocentral seta is missing, leaving a gap between first and third setae (H. omissa, H. confinis, H. triangularis, H. destituta, H. crypta, H. biseriata, H. bivittata). Houghia bivittata shares with H. biseriata the presence of a pair of wide dark scutal stripes (Figs. 13a, 14a) as a result of dark brown tomentosity between the two longitudinal dark scutal stripes on each side. The fronto-orbital plate is gold tomentose from vertex to antennal bases (Fig. 14b). Description. Male. Antenna black. When viewed in profile, antenna arises, distinctly above middle of eye. Length of first flagellomere extending to facial margin. Facial ridge bare except for a few (usually 3–5) decumbent small setae above vibrissa. Palpus pale, usually distinctly yellowish. Postgena behind postoccipital row, above level of lower facial margin, without black setae or with a small patch of few black setae. Parafacial silver. Colour of fronto-orbital plate pale brassy to gold on its entire length from vertex to base of antenna (more than 50% coverage). Surface of fronto-orbital plate almost bare. Ocellar triangle, when viewed from above appearing concolorous with frontal vitta. Diameter of anterior ocellus less than diameter of base of adjacent ocellar seta. Ocellar setae arising behind anterior ocellus but closer to anterior than to posterior ocelli. Eye bare. Postpronotum restricted to the three main postpronotal setae. Dark stripes on either side of dorsocentral row of setae, separated from one another with brownish or gray tomentosity, appearing paler than stripes, but darker than remainder of scutum. Median and lateral stripes on either side of scutum united to one another posteriorly. Postsutural dorsocentral setae 3, with a gap between first and third seta. Anterior quadrant of anepisternum covered with short setae except for usually 3 to 5 distinctly larger setae. Katepisternum bearing three setae, the middle one always the smallest. Vein R1 bare dorsally. Legs ranging from reddish brown to yellow tinged but overall dark. Coxae dark usually concolourous with remainder of leg. Ground colour of dorsal surface of abdomen dark to black. Ground colour of ventral surface of abdomen entirely black. Sex patches present on tergites 4 and 5. Ground colour of sex patches shiny black. Terminalia: surstylus equilaterally oblong shaped, posterodorsal half haired, apex bearing many stout apical spines, tip with strong inwardly apical curve when viewed dorsally. Cerci sharply pointed or distinctly tapered, apex with blunt, hooked tip, ventral surface haired, separation between cerci deep narrow V shape, up to 85% as long as surstylus. Lobe of sternite 5 large and rounded apically, inner margin covered in dense tomentosity appearing darker than surrounding cuticle, internal edge inwardly curved, single long apical seta emanating from apex, lobe size large and rounded apically. HOUGHIA OF ACG, COSTA RICA Zootaxa 3858 (1) © 2014 Magnolia Press · 37 Hosts. Houghia bivittata has been reared 25 times, from the combination of Euptychia westwoodi Butler and Euptychia mollis Staudinger (Satyrinae, Nymphalidae) feeding on the foliage of rain forest Selaginella arthritica Alston (Selaginaceae), in a sample of 351 caterpillars of these two species of butterflies (whose caterpillars are currently indistinguishable from each other). Holotype. ♂, CNC. Type locality: Costa Rica, Area de Conservación Guanacaste, Prov. Alajuela, Sector Rincon Rain Forest, Quebrada Escondida (10.89928°, -85.27486°), 420 m, 01/04/2011, Jose Perez, DHJPAR0042656. Paratypes. 9 ♂, 11 ♀ (CNC) Costa Rica, Prov. Alajuela and Guanacaste, ACG database codes: DHJPAR0048576, DHJPAR0048628, DHJPAR0048577, DHJPAR0046667, DHJPAR0046648, DHJPAR0048575, DHJPAR0046625, DHJPAR0046614, DHJPAR0046668, DHJPAR0046631, DHJPAR0046633, DHJPAR0046634, DHJPAR0048548, DHJPAR0048547, DHJPAR0048574, DHJPAR0046644, DHJPAR0046663, DHJPAR0048640. Etymology. From the Latin adverb,”bis”, “bi-” meaning two, and the adjective “vittatus” derived from “vitta”, meaning ribbon or stripe, in reference to the pair of broad dark stripes on the scutum. Distribution. Costa Rica, ACG, Prov. Alajuela & Guanacaste, rain forest and dry forest, 400–710 m elevation. FIGURE 14. General morphology of Houghia bivittata sp. nov. a. Dorsal habitus. b. Frontal view of head. c. Lateral habitus. d. Posterior view of male terminalia. e. Lateral view of male terminalia. f. Ventral view of sternite 5 + segments 7+8. 38 · Zootaxa 3858 (1) © 2014 Magnolia Press FLEMING ET AL. Houghia blancoi Fleming & Wood, sp. nov. Fig. 15 a–f Diagnosis. Houghia blancoi is a member of a group of five sibling species with few distinctive features except in the male terminalia and the extent of the gold tomentosity on the fronto-orbital plate; these are H. blancoi, H. romeroae, H. chavarriae, H. marini, and H. matarritai. The eye is bare and the antenna is entirely black. The ground colour of the abdomen is black, and sex patches are confined to tergites 4 and 5. In H. blancoi the gold tomentosity is confined to the vertex, not extending forward beyond the upper reclinate orbital seta (Fig. 15b) and the apex of the first flagellomere is shorter than the face by half the length of the pedicel. FIGURE 15. General morphology of Houghia blancoi sp. nov. a. Dorsal habitus. b. Frontal view of head. c. Lateral habitus. d. Posterior view of male terminalia. e. Lateral view of male terminalia. f. Ventral view of sternite 5 + segments 7+8. Description. Male. Antenna black. When viewed in profile, antenna arises distinctly above middle of eye. Length of first flagellomere almost extending to facial margin, (usually shorter by less than length of pedicel). Facial ridge bare except for a few (usually 3–5) decumbent small setae above vibrissa. Palpus pale, usually distinctly yellowish. Postgena behind postoccipital row, above level of lower facial margin, with a small patch of few black setae. Parafacial silver. Two possible forms exist, colour of fronto-orbital plate pale brassy to gold on its entire length from vertex to base of antenna (more than 50% coverage) or gold only at vertex, adjacent to ocellar triangle, the remainder silver (up to 25% coverage). Surface of fronto-orbital plate almost bare. Ocellar triangle, when viewed from above appearing to be notched anteriorly. Diameter of anterior ocellus equal to, or greater than, HOUGHIA OF ACG, COSTA RICA Zootaxa 3858 (1) © 2014 Magnolia Press · 39 diameter of base of adjacent ocellar seta or less than diameter of base of adjacent ocellar seta. Ocellar setae arising beside, or slightly in front of, anterior ocellus. Eye bare. Postpronotum with 4 or 5 postpronotal setae. Dark stripes on either side of dorsocentral row of setae separated from one another by yellow tomentosity. Median and lateral stripes on either side of scutum separate from each other posteriorly. Postsutural dorsocentral setae 4. Anterior quadrant of anepisternum covered with short setae except for usually 3 to 5 distinctly larger setae. Katepisternum with three setae, the middle one always the smallest. Vein R1 bare dorsally. Legs ranging from reddish brown to yellow tinged but overall dark. Coxae dark usually concolourous with remainder of leg. Ground colour of dorsal surface of abdomen dark to black. Ground colour of ventral surface of abdomen entirely black. Sex patches present on tergites 4 and 5. Ground colour of sex patches shiny black. Terminalia: surstylus bayonet shaped, posterodorsal half bare, apex bearing many stout apical spines, tip with light inwardly apical curve when viewed dorsally. Cerci rounded, apex with blunt, hooked tip, ventral surface haired, separation between cerci straight, about as long as surstylus. Lobe of sternite 5 small and pointed apically, inner margin covered in dense tomentosity appearing darker than surrounding cuticle, internal edge inwardly curved, multiple apical setae emanating from lobe. Hosts. Houghia blancoi has been reared 14 times from caterpillars of Trogoptera salvita (Mimallonidae). This widely-distributed caterpillar has been reared 552+ times in both dry forest and rain forest, however to date Houghia blancoi has only been reared from the dry forest. This fly was reared from no other species of caterpillar with the exception of one singleton fly barcode: DHJPAR0008143, which was reared from a megalopygid, Norape Janzen03. Holotype. ♂, CNC. Type locality Costa Rica, Area de Conservación Guanacaste, Prov. Guanacaste, Sector Horizontes, Vado Esperanza (10.78938°, -85.55098°), 85 m, 09/01/2007, Johan Vargas, DHJPAR0021839. Paratypes. 14 ♂, 17 ♀ (CNC) Costa Rica, Prov. Guanacaste, ACG database codes: DHJPAR0008136, DHJPAR0008139, DHJPAR0008143, DHJPAR0008138, DHJPAR0008137, DHJPAR0008839, DHJPAR0011508, DHJPAR0011530, DHJPAR0008121, DHJPAR0008111, DHJPAR0008134, DHJPAR0008135, DHJPAR0011490, DHJPAR0011529, DHJPAR0008133, 91-SRNP-2359, 90-SRNP-1732, 07-SRNP-14997, 90-SRNP-1730, 90SRNP-1775, 05-SRNP-65188. Etymology. Houghia blancoi is dedicated to Roger Blanco of Area de Conservación Guanacaste, cocoordinator of ACG Research and Subdirector for ACG Area Silvestre Protegido, in recognition of his decades of protection of the forests in which this tachinid lives. Distribution. Costa Rica, ACG, Prov. Guanacaste, dry forest, 85–380 m elevation. Houghia brevipilosa Fleming & Wood, sp. nov. Figs. 6 b, 16 a–f Diagnosis. One of a group of three species with long haired eyes, this species, and H. longipilosa have a predominantly silver head (entirely gold tomentose in H. aurifera). In H. brevipilosa only the upper part of the frons is gold tomentose, and the apical dark bands on abdominal tergites 3 to 5 are grayish brown rather than black (as in H. longipilosa), and therefore not as distinctly differentiated. Description. Male. Antenna black. When viewed in profile, antenna arises distinctly above middle of eye. Length of first flagellomere shorter than facial margin, (usually shorter by at least the length of the pedicel). Facial ridge bare except for a few (usually 3–5) decumbent small setae above vibrissa. Palpus pale, usually distinctly yellowish. Postgena behind postoccipital row, above level of lower facial margin, extensively covered with setae, and fine hairs. Parafacial silver. Colour of fronto-orbital plate gold on upper half, from vertex to midway down face (up to 50% coverage). Surface of fronto-orbital plate covered with small recumbent hairs, especially near margin of eye. Ocellar triangle, when viewed from above appearing to be notched anteriorly. Diameter of anterior ocellus equal to, or greater than, diameter of base of adjacent ocellar seta. Ocellar setae arising beside, or slightly in front of, anterior ocellus. Eye with conspicuous hairs. Postpronotum with 4 or 5 postpronotal setae. Dark stripes on either side of dorsocentral row of setae separated from one another by yellow tomentosity. Median and lateral stripes on either side of scutum separate from each other posteriorly. Postsutural dorsocentral setae 4. Anterior quadrant of anepisternum covered with long setae, none distinctly larger than the others or forming a row. Katepisternum with three setae, the middle one always the smallest. Vein R1 bare dorsally. Legs ranging from reddish brown to yellow tinged but overall dark. Coxae dark usually concolourous with remainder of leg. Ground 40 · Zootaxa 3858 (1) © 2014 Magnolia Press FLEMING ET AL. colour of dorsal surface of abdominal tergite 3 and in some specimens parts of other tergites as well, dark medially but reddish laterally. Ground colour of ventral surface of abdomen partially or entirely reddish or yellowish. Sex patches present on tergites 4 and 5. Ground colour of sex patches light coloured or pale. Terminalia: surstylus bayonet shaped, posterodorsal half bare, apex bearing few short apical spines, tip with light inwardly apical curve when viewed dorsally. Cerci rounded, apex with blunt, hooked tip, ventral surface haired, separation between cerci forms a deep narrow V shape, up to 85% as long as surstylus. Lobe of sternite 5 small and pointed apically, inner margin covered in dense tomentosity appearing darker than surrounding cuticle, internal edge inwardly curved, apical seta absent. FIGURE 16. General morphology of Houghia brevipilosa sp. nov. a. Dorsal habitus. b. Frontal view of head. c. Lateral habitus. d. Posterior view of male terminalia. e. Lateral view of male terminalia. f. Ventral view of sternite 5 + segments 7+8. Hosts. Houghia brevipilosa has been reared 130 times, from 6300+ wild-caught dry forest caterpillars of eudamine Hesperiidae, of 19 species in 9 genera, but primarily from Urbanus, Astraptes and Epargyreus. It has been reared only from this group of ACG caterpillars. Holotype. ♂, CNC. Type locality: Costa Rica, Area de Conservación Guanacaste, Prov. Guanacaste, Sector Santa Rosa, Vado Cuajiniquil (10.94041°, -85.68043°), 275 m, 02/15/2006, Freddy Quesada, DHJPAR0006994. Paratypes. 34 ♂, 16 ♀ (CNC) Costa Rica, Prov. Guanacaste, ACG database codes: DHJPAR0008201, DHJPAR0008203, DHJPAR0008200, DHJPAR0008205, DHJPAR0008209, DHJPAR0008208, DHJPAR0027950, DHJPAR0027951, DHJPAR0008174, DHJPAR0008179, DHJPAR0008196, DHJPAR0008199, DHJPAR0008216, HOUGHIA OF ACG, COSTA RICA Zootaxa 3858 (1) © 2014 Magnolia Press · 41 DHJPAR0008217, DHJPAR0008549, DHJPAR0008210, DHJPAR0008220, DHJPAR0008223, DHJPAR0008224, DHJPAR0008231, DHJPAR0008235, DHJPAR0007025, DHJPAR0034359, DHJPAR0006990, DHJPAR0017193, DHJPAR0006992, DHJPAR0007091, DHJPAR0027949, DHJPAR0006968, DHJPAR0037435, DHJPAR0037438, DHJPAR0008186, DHJPAR0008191, DHJPAR0008177, DHJPAR0008182, DHJPAR0008185, DHJPAR0008180, DHJPAR0019564, DHJPAR0008192, DHJPAR0008189, DHJPAR0008176, DHJPAR0008178, DHJPAR0008183, DHJPAR0008197, DHJPAR0008194, DHJPAR0008226, DHJPAR0008175, DHJPAR0008214, DHJPAR0008190, DHJPAR0008228, 09-SRNP-12053, 07-SRNP-20253, 95-SRNP-5156, 94-SRNP-10107, 00-SRNP-6265. Etymology. From the Latin adjective, “brevis”, meaning short, and the adjective “pilosus”, covered with hair, in reference to the conspicuous eye hairs that appear to be slightly shorter than those of H. aurifera and H. longipilosa, described below. Distribution. Costa Rica, ACG, Prov. Guanacaste, dry forest and rain–dry forest ecotones, 8–820 m elevation. Houghia chavarriae Fleming & Wood, sp. nov. Figs. 2 b, 17 a–f Diagnosis. Houghia chavarriae is a member of the H. blancoi group of sibling species (H. blancoi, H. romeroae, H. chavarriae, H. marini, and H. matarritai), with few distinctive features except in the male terminalia and in the extent of gold tomentosity on the fronto-orbital plate. The eye is bare and the antenna is entirely black. The ground colour of the abdomen is black, and sex patches are confined to tergites 4 and 5. The gold tomentosity of the frons of H. chavarriae extends from the vertex to the level of the anterior proclinate orbital setae (Fig. 17b). In lateral view (Fig. 17e) the surstylus is obovate, or widest in the mid-region rather than straight-sided, as in H. spathulata (Fig. 43e) while its posterior margin is rounded posteriorly, compared to the concave posterior margin of H. matarritai (Fig. 34e). Description. Male. Antenna black. When viewed in profile, antenna arises approximately at level of middle of eye. Length of first flagellomere shorter than facial margin, (usually shorter by at least the length of the pedicel). Facial ridge bare except for a few (usually 3–5) decumbent small setae above vibrissa. Palpus pale, usually distinctly yellowish. Postgena behind postoccipital row, above level of lower facial margin, with a small patch of few black setae. Parafacial silver. Colour of fronto-orbital plate pale brassy to gold on its entire length from vertex to base of antenna (more than 50% coverage). Surface of fronto-orbital plate covered with small recumbent hairs, especially near margin of eye. Ocellar triangle, when viewed from above appearing rounded anteriorly. Diameter of anterior ocellus equal to, or greater than, diameter of base of adjacent ocellar seta. Ocellar setae arising beside, or slightly in front of, anterior ocellus. Eye with inconspicuous hairs. Postpronotum with 4 or 5 postpronotal setae. Dark stripes on either side of dorsocentral row of setae separated from one another by yellow tomentosity. Median and lateral stripes on either side of scutum separate from each other posteriorly. Postsutural dorsocentral setae 4. Anterior quadrant of anepisternum covered with short setae except for usually 3 to 5 distinctly larger setae. Katepisternum with three setae, the middle one always extremely reduced, might be absent in some individuals. Vein R1 bare dorsally. Legs ranging from reddish brown to yellow tinged but overall dark. Coxae dark usually concolourous with remainder of leg. Ground colour of dorsal surface of abdomen dark to black. Ground colour of ventral surface of abdomen partially or entirely reddish or yellowish. Sex patches present on tergites 4 and 5. Ground colour of sex patches light coloured or pale. Terminalia: surstylus equilaterally oblong shaped, posterodorsal half bare, apex bearing many stout apical spines, tip with light inwardly apical curve when viewed dorsally. Cerci rounded, apex with blunt, hooked tip, ventral surface haired, separation between cerci straight, up to 85% as long as surstylus. Lobe of sternite 5 small and pointed apically, inner margin covered in dense tomentosity appearing darker than surrounding cuticle, internal edge inwardly curved, single long apical seta emanating from apex. Hosts. Houghia chavarriae has been reared 14 times, from 10000+ wild-caught dry forest caterpillars of macroglossine Sphingidae, of 6 species in 4 genera, but primarily from Eupyrrhoglossum sagra (Poey) and Nyceryx coffaeae Walker. It has been reared only from this group of ACG caterpillars; all rearings are from dry forest, even though the host caterpillars occur in both rain forest and dry forest. Holotype. ♂, CNC. Type locality: Costa Rica, Area de Conservación Guanacaste, Prov. Guanacaste, Sector Santa Rosa, Bosque San Emilio (10.84389°, -85.61384°), 300 m, 06/06/2005, Ruth Franco, DHJPAR0008127. 42 · Zootaxa 3858 (1) © 2014 Magnolia Press FLEMING ET AL. FIGURE 17. General morphology of Houghia chavarriae sp. nov. a. Dorsal habitus. b. Frontal view of head. c. Lateral habitus. d. Posterior view of male terminalia. e. Lateral view of male terminalia. f. Ventral view of sternite 5 + segments 7+8. Paratypes. 82 ♂, 89 ♀ (CNC) Costa Rica, Guanacaste, ACG database codes: DHJPAR0029781, DHJPAR0020943, DHJPAR0008115, DHJPAR0008838, DHJPAR0008544, DHJPAR0008140, DHJPAR0008114, DHJPAR0008113, DHJPAR0008129, DHJPAR0008112, DHJPAR0008162, 00-SRNP-2771, 01-SRNP-16535, 05SRNP-15809, 08-SRNP-14718, 84-SRNP-2119, 92-SRNP-3614, 81-SRNP-913, 96-SRNP-7189, 81-SRNP-574, 81-SRNP-564, 07-SRNP-58359, 01-SRNP-17033, 01-SRNP-16458. Etymology. Houghia chavarriae is dedicated to Srta. Maria Marta Chavarria of Area de Conservación Guanacaste, coordinator of ACG Research and of the Marine Biosensitivity Program, in recognition of her decades of protection of the forests in which this tachinid lives. Distribution. Costa Rica, ACG, Prov. Guanacaste, dry forest, 95–395 m elevation. Houghia confinis Fleming & Wood, sp. nov. Figs. 10 d, 18 a–f Diagnosis. One of a group of seven closely related species in ACG, and many more in Central and South America, the H. crypta group, in which the second postsutural dorsocentral seta is missing, leaving a gap between first and third setae (H. omissa, H. confinis, H. triangularis, H. destituta, H. crypta, H. biseriata, H. bivittata). Houghia HOUGHIA OF ACG, COSTA RICA Zootaxa 3858 (1) © 2014 Magnolia Press · 43 confinis may be distinguished from the other six by the converging dark scutal stripes; the outer stripe on each side curves medially toward the second last postsutural dorsocentral seta to meet or nearly meet the inner stripe (Fig. 18a). FIGURE 18. General morphology of Houghia confinis sp. nov. a. Dorsal habitus. b. Frontal view of head. c. Lateral habitus. d. Posterior view of male terminalia. e. Lateral view of male terminalia. f. Ventral view of sternite 5 + segments 7+8. Description. Male. Antenna black. When viewed in profile, antenna arises, approximately at level of middle of eye. Length of first flagellomere almost extending to facial margin, (usually shorter by less than length of pedicel). Facial ridge bare except for a few (usually 3–5) decumbent small setae above vibrissa. Palpus pale, usually distinctly yellowish. Postgena behind postoccipital row, above level of lower facial margin, with a small patch of few black setae. Parafacial silver. Colour of fronto-orbital plate gold only at vertex, adjacent to ocellar triangle, the remainder silver (up to 25% coverage). Surface of fronto-orbital plate almost bare. Ocellar triangle, when viewed from above appearing to be sharply pointed anteriorly. Diameter of anterior ocellus equal to, or greater than, diameter of base of adjacent ocellar seta. Ocellar setae arising beside, or slightly in front of, anterior ocellus. Eye bare. Postpronotum restricted to the three main postpronotal setae. Dark stripes on either side of dorsocentral row of setae, separated from one another with brownish or gray tomentosity, appearing paler than stripes, but darker than remainder of scutum. Median and lateral stripes on either side of scutum united to one another posteriorly. Postsutural dorsocentral setae 3, with a gap between first and third seta. Anterior quadrant of anepisternum covered with long setae, none distinctly larger than the others or forming a row. Katepisternum bearing three setae, the middle one always the smallest. Vein R1 bare dorsally. Legs ranging from reddish brown to yellow tinged but 44 · Zootaxa 3858 (1) © 2014 Magnolia Press FLEMING ET AL. overall dark. Coxae dark usually concolourous with remainder of leg. Ground colour of dorsal surface of abdomen dark to black. Ground colour of ventral surface of abdomen entirely black. Sex patches present on tergites 4 and 5. Ground colour of sex patches shiny black. Terminalia: surstylus equilaterally oblong shaped, posterodorsal half haired, apex bearing few short apical spines, tip with strong inwardly apical curve when viewed dorsally. Cerci rounded, apex with blunt, hooked tip, ventral surface bare, separation between cerci straight, about as long as surstylus. Lobe of sternite 5 small and pointed apically, inner margin covered in dense tomentosity appearing darker than surrounding cuticle, internal edge inwardly curved, apical seta absent. Hosts. Houghia confinis has been reared 17 times, and entirely from a sample of 3000+ grass-eating butterflies (Nymphalidae) belonging to seven genera that eat only Poaceae, except for a single additional rearing from a wildcaught pupa of an unknown (and unknowable) species of hesperiine skipper butterfly (Hesperiidae). However, there also remains the possibility that what was identified as a hesperiine pupa might have been actually a nymphalid butterfly pupa. Holotype. ♂, CNC. Type locality: Costa Rica, Area de Conservación Guanacaste, Prov. Guanacaste, Sector Pitilla, Sendero Laguna (10.98880°, -85.42336°), 680 m, 09/28/2011, Freddy Quesada, DHJPAR0046632. Paratypes. 8 ♂, 10 ♀ (CNC) Costa Rica, Guanacaste, ACG database codes: DHJPAR0048658, DHJPAR0008786, DHJPAR0045641, DHJPAR0021835, DHJPAR0008790, DHJPAR0046618, DHJPAR0046568, DHJPAR0048570, DHJPAR0044896, DHJPAR0046624, DHJPAR0048563, DHJPAR0008784, DHJPAR0021968, DHJPAR0008529, DHJPAR0048569, DHJPAR0045595, DHJPAR0048571. Etymology. From the Latin adjective, “confinis”, meaning adjacent, in reference to the convergence of the pair of dark stripes on each side of the scutum that unite or converge toward one another at the level of the third postsutural dorsocentral seta. Distribution. Costa Rica, ACG, Prov. Alajuela & Guanacaste, rain forest and dry forest, 160–680 m elevation. Houghia crypta (Townsend, 1934 402) Fig. 19 a–f Carceliocephala crypta Townsend, 1934: 402. Holotype female (USNM) [examined by DMW]. Type locality: Brazil, Para, Río Tapajós, Bôa Vista. Diagnosis. One of a group of seven sibling species in ACG, and many more in Central and South America, that we have called the H. crypta group (Townsend’s monotypic genus Carceliocephala), in which the second postsutural dorsocentral seta is missing, leaving a gap between first and third setae (H. omissa, H. confinis, H. triangularis, H. destituta, H. crypta, H. biseriata, H. bivittata). We interpret this species as conspecific with the holotype of Houghia crypta (Townsend) described from Brazil. It may be distinguished from the remaining 6 species by the presence of a sex patch on tergite 5 (absent in H. omissa), two distinctly separated stripes on either side of the scutum (these stripes united by dark tomentosity in H. bivittata and H. bistrigata thus appearing as a single broad stripe on each side, or partially united posteriorly, as in H. confinis), by the truncated or notched ocellar triangle, by the long dagger-like surstylus, and by the shape of the cerci (explained in couplet 17). However, the holotype has not been dissected, so comparisons of the terminalia with those of the holotype have not been made. Description. Male. Antenna black. When viewed in profile, antenna arises approximately at level of middle of eye. Length of first flagellomere shorter than facial margin, (usually shorter by at least the length of the pedicel). Facial ridge bare except for a few (usually 3–5) decumbent small setae above vibrissa. Palpus pale, usually distinctly yellowish. Postgena behind postoccipital row, above level of lower facial margin, with a small patch of few black setae. Parafacial silver. Colour of fronto-orbital plate gold on upper half, from vertex to midway down face (up to 50% coverage). Surface of fronto-orbital plate almost bare. Ocellar triangle, when viewed from above appearing to be notched anteriorly. Diameter of anterior ocellus equal to, or greater than, diameter of base of adjacent ocellar seta. Ocellar setae arising beside, or slightly in front of, anterior ocellus. Eye bare. Postpronotum with 4 or 5 postpronotal setae. Dark stripes on either side of dorsocentral row of setae, separated from one another by yellow tomentosity. Median and lateral stripes on either side of scutum separate from each other posteriorly. Postsutural dorsocentral setae 3, with a gap between first and third seta. Anterior quadrant of anepisternum covered with short setae except for usually 3 to 5 distinctly larger setae. Katepisternum bearing three setae, the middle one HOUGHIA OF ACG, COSTA RICA Zootaxa 3858 (1) © 2014 Magnolia Press · 45 always the smallest. Vein R1 bare dorsally. Legs ranging from reddish brown to yellow tinged but overall dark. Coxae dark usually concolourous with remainder of leg. Ground colour of dorsal surface of abdomen dark to black. Ground colour of ventral surface of abdomen entirely black. Sex patches present on tergites 4 and 5. Ground colour of sex patches shiny black. Terminalia: surstylus equilaterally long dagger-like, posterodorsal half bare, apex bearing many stout apical spines, tip with strong inwardly apical curve when viewed dorsally. Cerci rounded, apex with blunt, hooked tip, ventral surface bare, separation between cerci deep narrow V shape, up to 85% as long as surstylus. Lobe of sternite 5 small and pointed apically, inner margin covered in dense tomentosity appearing darker than surrounding cuticle, internal edge inwardly curved, apical seta absent. Hosts. Houghia crypta has been reared 18 times, from a sample of 896 caterpillars of Callopistria floridensis (Guenée) (Noctuidae) feeding on seven species of dry forest and rain forest ferns in 5 families. Type Material Examined. ♂, (USNM): BV.15.VII// Type// Carceliocephala/crypta TT/ ♂ /DetCHTT// Type/ Ht ♂ [examined by DMW]. Other Material Examined. 7 ♂, 7 ♀ (CNC) Costa Rica, Prov. Alajuela and Guanacaste, ACG database codes: DHJPAR0016659, DHJPAR0008781, DHJPAR0016709, DHJPAR0046649, DHJPAR0016658, DHJPAR0016716, DHJPAR0008789, DHJPAR0046630, DHJPAR0016717, DHJPAR0042606, DHJPAR0016657, DHJPAR0016708, DHJPAR0046643, DHJPAR0046647, 1 ♀ Costa Rica, H[acien]da Chilamate Selva Verde lodge, [Sarapiqui, Prov. Heredia] 21-22.IX.89 M. Polak. Distribution. Costa Rica, ACG, Prov. Alajuela & Guanacaste, rain forest and dry forest, 95–580 m elevation. FIGURE 19. General morphology of Houghia crypta sp. nov. a. Dorsal habitus. b. Frontal view of head. c. Lateral habitus. d. Posterior view of male terminalia. e. Lateral view of male terminalia. f. Ventral view of sternite 5 + segments 7+8. 46 · Zootaxa 3858 (1) © 2014 Magnolia Press FLEMING ET AL. Houghia delospilota Fleming & Wood, sp. nov. Fig. 20 a–f Diagnosis. One of 5 species with reddish or yellowish abdominal tergites 3 and 4, The sex patches of this species are restricted to tergites 4 and 5; these consist of black appressed hairs on either a pale reddish ground colour or on a spot with some darker pigment beneath, but otherwise the ventral surfaces of the tergites are pale. The frontoorbital plate is mostly silver tomentose, with gold tomentosity and short recumbent hairs, extending from vertex only to the level of the anterior fronto-orbital setae (Fig. 20b). FIGURE 20. General morphology of Houghia delospilota sp. nov. a. Dorsal habitus. b. Frontal view of head. c. Lateral habitus. d. Posterior view of male terminalia. e. Lateral view of male terminalia. f. Ventral view of sternite 5 + segments 7+8. Description. Male. Antenna black. When viewed in profile, antenna arises distinctly above middle of eye. Length of first flagellomere shorter than facial margin, (usually shorter by at least the length of the pedicel). Facial ridge bare except for a few (usually 3–5) decumbent small setae above vibrissa. Palpus pale, usually distinctly yellowish. Postgena behind postoccipital row, above level of lower facial margin, with a small patch of few black setae. Parafacial silver. Colour of fronto-orbital plate gold on upper half, from vertex to midway down face (up to 50% coverage). Surface of fronto-orbital plate covered with small recumbent hairs, especially near margin of eye. Ocellar triangle, when viewed from above appearing to be notched anteriorly. Diameter of anterior ocellus equal to, or greater than, diameter of base of adjacent ocellar seta. Ocellar setae arising beside, or slightly in front of, anterior ocellus. Eye bare. Postpronotum with 4 or 5 postpronotal setae. Dark stripes on either side of dorsocentral row of HOUGHIA OF ACG, COSTA RICA Zootaxa 3858 (1) © 2014 Magnolia Press · 47 setae separated from one another with brownish or gray tomentosity, appearing paler than stripes, but darker than remainder of scutum. Median and lateral stripes on either side of scutum separate from each other posteriorly. Postsutural dorsocentral setae 4. Anterior quadrant of anepisternum covered with short setae except for usually 3 to 5 distinctly larger setae. Katepisternum with three setae, the middle one always the smallest. Vein R1 bare dorsally. Legs ranging from reddish brown to yellow tinged but overall dark. Coxae dark usually concolourous with remainder of leg. Ground colour of dorsal surface of abdominal tergite 3 and in some specimens parts of other tergites as well, dark medially but reddish laterally. Ground colour of ventral surface of abdomen partially or entirely reddish or yellowish. Sex patches present on tergites 4 and 5. Ground colour of sex patches not distinctly shiny. Terminalia: surstylus bayonet shaped, posterodorsal half bare, apex bearing many stout apical spines, tip with strong inwardly apical curve when viewed dorsally. Cerci rounded, apex with blunt, hooked tip, ventral surface haired, separation between cerci deep narrow V shape, up to 85% as long as surstylus. Lobe of sternite 5 large and rounded apically, inner margin covered in dense tomentosity appearing darker than surrounding cuticle, internal edge inwardly curved, apical seta absent. Hosts. Houghia delospilota has been reared 13 times, from four species of dry forest Nystalea Guenée (Notodontidae), N. collarisDHJ13 (1X), N. guzmani Schaus (3X), N. morona (Druce) (8X), and N. multiplex Dognin (1X), among a sample of 2500+ Nystalea caterpillars. Holotype. ♂, CNC. Type locality: Costa Rica, Area de Conservación Guanacaste, Prov. Guanacaste, Sector El Hacha, Sendero Bejuquilla (11.03004°, -85.52699°), 280 m, 02/03/2000, gusaneros, DHJPAR0008837. Paratypes. 38 ♂, 49 ♀ (CNC) Costa Rica, Prov. Guanacaste, ACG database codes: DHJPAR0008828, DHJPAR0048672, DHJPAR0008834, DHJPAR0029712, DHJPAR0008826, DHJPAR0008833, DHJPAR0008831, DHJPAR0008827, DHJPAR0008832, DHJPAR0008845, DHJPAR0008829, DHJPAR0008836, DHJPAR0011472, 00-SRNP-2271, 12-SRNP-20327, 99-SRNP-16134, 03-SRNP-19519, 02-SRNP-29961, 00-SRNP-2110, 98SRNP-14306, 02-SRNP-29968, 05-SRNP-47671. Etymology. From Greek, delos, evident, and spilotos, spotted, here intended as an adjective to describe the conspicuous sex patches on tergites 4 and 5 (use of the name as an adjective was discussed by Orr & Fliedner 2011). Distribution. Costa Rica, ACG, Prov. Guanacaste, rain forest and dry forest, 220–550 m elevation. Houghia destituta Fleming & Wood, sp. nov. Fig. 21 a–f Diagnosis. One of a group of seven closely related species in ACG, and many more in Central and South America, the H. crypta group, in which the second postsutural dorsocentral seta is missing, leaving a gap between first and third setae (H. omissa, H. confinis, H. triangularis, H. destituta, H. crypta, H. biseriata, H. bivittata) (Figs. 7b, 21d) (a characteristic of the H. crypta group). Differentiates itself from the rest of the crypta group by the depth of the dorsal notch (Fig. 21d) on the cerci in posterior view, that appears to be as high as the apical portion (Fig. 21d). Description. Male. Antenna black. When viewed in profile, antenna arises approximately at level of middle of eye. Length of first flagellomere almost extending to facial margin (usually shorter by less than length of pedicel). Facial ridge bare except for a few (usually 3–5) decumbent small setae above vibrissa. Palpus pale, usually distinctly yellowish. Postgena behind postoccipital row, above level of lower facial margin, with a small patch of few black setae. Parafacial silver. Colour of fronto-orbital plate gold on upper half, from vertex to midway down face (up to 50% coverage). Surface of fronto-orbital plate almost bare. Ocellar triangle, when viewed from above appearing to be sharply pointed anteriorly. Diameter of anterior ocellus less than diameter of base of adjacent ocellar seta. Ocellar setae arising beside, or slightly in front of, anterior ocellus. Eye bare. Postpronotum restricted to the three main postpronotal setae. Dark stripes on either side of dorsocentral row of setae, separated from one another by yellow tomentosity. Median and lateral stripes on either side of scutum united to one another posteriorly. Postsutural dorsocentral setae 3, with a gap between first and third seta. Anterior quadrant of anepisternum covered with short setae except for usually 3 to 5 distinctly larger setae. Katepisternum bearing three setae, the middle one always the smallest. Vein R1 bare dorsally. Legs ranging from reddish brown to yellow tinged but overall dark. Coxae distinctly yellow tinged, usually contrasting with darker remainder of legs. Ground colour of dorsal surface of abdomen dark to black. Ground colour of ventral surface of abdomen entirely black. Sex 48 · Zootaxa 3858 (1) © 2014 Magnolia Press FLEMING ET AL. patches present on tergites 4 and 5. Ground colour of sex patches shiny black. Terminalia: surstylus bayonet shaped, posterodorsal half bare, apex bearing apical spines along lower half only, tip with strong inwardly apical curve when viewed dorsally. Cerci rounded, apex with blunt, hooked tip, ventral surface bare, separation between cerci straight, up to 85% as long as surstylus. Lobe of sternite 5 small and pointed apically, inner margin covered in dense tomentosity appearing darker than surrounding cuticle, internal edge inwardly curved, single long apical seta emanating from apex. Hosts. Houghia destituta has been reared 9 times, from a sample of 284 caterpillars of the rain forest Antiblemma ceras (Druce) (Noctuidae) feeding on Conostegia xalapensis (Bonpl.) (Melastomataceae). Holotype. ♂, CNC. Type locality: Costa Rica, Area de Conservación Guanacaste, Prov. Alajuela, Sector Rincon Rain Forest, Sendero Venado (10.89678°, -85.27001°), 420 m, 11/17/2008, Anabelle Cordoba, DHJPAR0030015. Paratypes. 3 ♂, 6 ♀ (CNC) Costa Rica, Prov. Alajuela and Guanacaste, ACG database codes: DHJPAR0042295, DHJPAR0007013, DHJPAR0008787, DHJPAR0030010, 00-SRNP-20618, DHJPAR0016245, DHJPAR0036549, 00-SRNP-20588. Etymology. From the Latin participle “destitutus”, derived from the verb “destituo” meaning deserted, devoid of, or lacking, referring to the paucity of distinguishing features. Distribution. Costa Rica, ACG, Prov. Alajuela & Guanacaste, rain forest and dry forest, 135–461 m elevation. FIGURE 21. General morphology of Houghia destituta sp. nov. a. Dorsal habitus. b. Frontal view of head. c. Lateral habitus. d. Posterior view of male terminalia. e. Lateral view of male terminalia. f. Ventral view of sternite 5 + segments 7+8. HOUGHIA OF ACG, COSTA RICA Zootaxa 3858 (1) © 2014 Magnolia Press · 49 Houghia fimbriata Fleming & Wood, sp. nov. Figs. 1 b, 3, 6 a, 7 a, 22 a–f Diagnosis. The most distinctive feature of this species, although difficult to see and easily overlooked, is the presence on the lower half of the facial ridge of a row of minute white hairs, best observed against a dark background (Fig. 3). Supporting character states are the orange base of the first flagellomere (also present in H. macilenta, H. nigrofemur, H. ochrofemur, and H. pallida) and the almost entirely silver and tomentose head. FIGURE 22. General morphology of Houghia fimbriata sp. nov. a. Dorsal habitus. b. Frontal view of head. c. Lateral habitus. d. Posterior view of male terminalia. e. Lateral view of male terminalia. f. Ventral view of sternite 5 + segments 7+8. Description. Male. Pedicel and upper portion of first flagellomere pale orange. When viewed in profile, antenna arises distinctly above middle of eye. Length of first flagellomere almost extending to facial margin (usually shorter by less than length of pedicel). Facial ridge with, in addition to the decumbent vibrissae, a row of minute, inconspicuous, hairs. Palpus pale, usually distinctly yellowish. Postgena behind postoccipital row, above level of lower facial margin, with a small patch of few black setae. Parafacial silver. Colour of fronto-orbital plate gold only at vertex, adjacent to ocellar triangle, the remainder silver (up to 25% coverage). Surface of fronto-orbital plate covered with small recumbent hairs, especially near margin of eye. Ocellar triangle viewed from above appearing to be sharply pointed anteriorly. Diameter of anterior ocellus equal to, or greater than, diameter of base of adjacent ocellar seta. Ocellar setae arising behind anterior ocellus but closer to anterior than to posterior ocelli. Eye bare. Postpronotum restricted to the three main postpronotal setae. Dark stripes on either side of dorsocentral 50 · Zootaxa 3858 (1) © 2014 Magnolia Press FLEMING ET AL. row of setae separated from one another by yellow tomentosity. Median and lateral stripes on either side of scutum separate from each other posteriorly. Postsutural dorsocentral setae 3. Anterior quadrant of anepisternum covered with long setae, none distinctly larger than the others or forming a row. Katepisternum with two setae. Vein R1 bare dorsally. Legs ranging from reddish brown to yellow tinged but overall dark. Coxae dark usually concolourous with remainder of leg. Ground colour of dorsal surface of abdominal tergite 3 and in some specimens parts of other tergites as well, dark medially but reddish laterally. Ground colour of ventral surface of abdomen partially or entirely reddish or yellowish. Sex patches present on tergite 3, as well as on tergites 4 and 5 (though sometimes small or absent on 5). Ground colour of sex patches either not distinctly shiny, or light coloured or pale. Terminalia: surstylus wedge shaped, posterodorsal half haired, apex bearing few short apical spines, tip straight. Cerci rounded, apex with blunt, hooked tip, ventral surface haired, separation between cerci straight, length of cerci up to 85% as long as surstylus. Lobe of sternite 5 large and rounded apically, inner margin covered in dense tomentosity appearing darker than surrounding cuticle, internal edge of lobe straight, with single long apical seta emanating from apex. Hosts. 666 wild-caught Zaretis ellops (Ménétriés) (Nymphalidae) caterpillars collected from ACG dry forest between 1987 and 2012 produced 60 rearings of this fly. It has been reared only from Zaretis ellops. Holotype. ♂, CNC. Type locality: Costa Rica, Area de Conservación Guanacaste, Prov. Guanacaste, Sector Cacao, Quebrada Heliconia (10.88585°, -85.49222°), 390 m, 12/25/2004, Manuel Pereira, DHJPAR0019206. Paratypes. 22 ♂, 21 ♀ (CNC) Costa Rica, Prov. Guanacaste, ACG database codes: DHJPAR0019224, DHJPAR0019205, DHJPAR0019218, DHJPAR0019216, DHJPAR0029805, DHJPAR0019221, DHJPAR0029802, DHJPAR0019215, DHJPAR0016048, DHJPAR0019609, DHJPAR0019213, DHJPAR0019969, DHJPAR0019209, DHJPAR0011475, DHJPAR0016742, DHJPAR0019207, DHJPAR0011478, DHJPAR0019204, DHJPAR0049537, DHJPAR0019220, DHJPAR0019203, DHJPAR0019222, DHJPAR0019210, DHJPAR0011477, DHJPAR0019211, DHJPAR0011511, DHJPAR0019219, DHJPAR0040986, DHJPAR0019217, DHJPAR0019208, DHJPAR0011476, DHJPAR0019223, DHJPAR0020922, DHJPAR0016647, DHJPAR0019214, DHJPAR0011474, DHJPAR0016683, DHJPAR0019212, 06-SRNP-47950, 97-SRNP-10010, 95-SRNP-8669. Etymology. From the Latin adjective, fimbriatus, meaning fringed, referring to the distinctive row of tiny hairs extending along the facial ridge. Similar hairs are present in other species, but they extend less than half way to the base of the antenna. Distribution. Costa Rica, ACG, Prov. Guanacaste, dry forest, 85–740 m elevation. Houghia gracilis Fleming & Wood, sp. nov. Fig. 23 a–f Diagnosis. One of a pair of sibling species, the other being H. graciloides, with orange ground colour on sides and ventral surface of tergites 3 and 4 and extensive but sparse sex patches on tergites 3 to 5; with ocellar setae arising distinctly behind anterior ocellus, and with a row of minute black hairs on lower 1/4 of facial ridge above the usual decumbent supravibrissal setae (H. fimbriata has minute whitish hairs on lower half of facial ridge; H. sexmaculata has sex patches on tergites 3 to 5 on a black ground colour, and several species have the ocellar setae arising behind the anterior ocellus). Eye apparently bare, even under high magnification. Description. Male. Antenna black. When viewed in profile, antenna arises approximately at level of middle of eye. Length of first flagellomere extending to facial margin. Facial ridge with, in addition to the decumbent vibrissae, a row of minute, inconspicuous, hairs. Palpus pale, usually distinctly yellowish. Postgena behind postoccipital row, above level of lower facial margin, with a small patch of few black setae. Parafacial silver. Colour of fronto-orbital plate pale brassy to gold on its entire length from vertex to base of antenna (more than 50% coverage). Surface of fronto-orbital plate almost bare. Ocellar triangle, when viewed from above appearing rounded anteriorly. Diameter of anterior ocellus equal to, or greater than, diameter of base of adjacent ocellar seta. Ocellar setae arising behind anterior ocellus but closer to anterior than to posterior ocelli. Eye bare. Postpronotum restricted to the three main postpronotal setae. Dark stripes on either side of dorsocentral row of setae separated from one another by yellow tomentosity. Median and lateral stripes on either side of scutum separate from each other posteriorly. Postsutural dorsocentral setae 4. Anterior quadrant of anepisternum covered with short setae except for usually 3 to 5 distinctly larger setae. Katepisternum with three setae, the middle one always the smallest. HOUGHIA OF ACG, COSTA RICA Zootaxa 3858 (1) © 2014 Magnolia Press · 51 Vein R1 bare dorsally. Legs ranging from reddish brown to yellow tinged but overall dark. Coxae distinctly yellow tinged, usually contrasting with darker remainder of legs. Ground colour of dorsal surface of abdominal tergite 3 and in some specimens parts of other tergites as well, dark medially but reddish laterally. Ground colour of ventral surface of abdomen partially or entirely reddish or yellowish. Sex patches present on tergite 3, as well as on tergites 4 and 5 (though sometimes small or absent on 5). Ground colour of sex patches not distinctly shiny. Terminalia: surstylus equilaterally oblong shaped, posterodorsal half haired, apex bearing many stout apical spines, tip with light inwardly apical curve when viewed dorsally. Cerci sharply pointed or distinctly tapered, apex with straight tip, ventral surface bare, separation between cerci straight, about as long as surstylus. Lobe of sternite 5 oversized and rounded apically, inner margin covered in dense tomentosity appearing darker than surrounding cuticle, internal edge straight, single long apical seta emanating from apex. FIGURE 23. General morphology of Houghia gracilis sp. nov. a. Dorsal habitus. b. Frontal view of head. c. Lateral habitus. d. Posterior view of male terminalia. e. Lateral view of male terminalia. f. Ventral view of sternite 5 + segments 7+8. Hosts. Houghia gracilis has been reared 43 times, from 27 species in 19 genera of primarily herbaceous monocot-eating caterpillars of Hesperiinae (Hesperiidae) in ACG dry forest, cloud forest, and rain forest, from a sample of 10000+ caterpillars. This long list of hosts overlaps with that of H. graciloides, but emphasizes grasses and other short-lived monocot leaves, instead of large and long-lived monocot leaves. Holotype. ♂, CNC. Type locality: Costa Rica, Area de Conservación Guanacaste, Prov. Alajuela, Sector San Cristobal, Sendero Perdido (10.8794°, -85.38607°), 620 m, 03/16/2004, Anabelle Cordoba, DHJPAR0008435. Paratypes. 67 ♂, 34 ♀ (CNC) Costa Rica, Prov. Guanacaste & Alajuela, ACG database codes: 52 · Zootaxa 3858 (1) © 2014 Magnolia Press FLEMING ET AL. DHJPAR0008405, DHJPAR0023638, DHJPAR0008388, DHJPAR0020981, DHJPAR0008428, DHJPAR0008410, DHJPAR0008434, DHJPAR0023687, DHJPAR0005478, DHJPAR0007105, DHJPAR0008426, DHJPAR0007004, DHJPAR0019553, DHJPAR0040663, DHJPAR0008389, DHJPAR0036598, DHJPAR0008395, DHJPAR0027864, DHJPAR0008435, DHJPAR0019556, DHJPAR0019555, DHJPAR0019739, DHJPAR0011062, DHJPAR0009995, DHJPAR0035680, DHJPAR0048669, DHJPAR0008736, DHJPAR0011501, DHJPAR0008384, DHJPAR0019658, DHJPAR0008401, DHJPAR0008383, DHJPAR0008420, DHJPAR0015016, DHJPAR0011502, DHJPAR0008402, DHJPAR0008377, DHJPAR0008378, DHJPAR0008429, DHJPAR0008407, DHJPAR0008427, DHJPAR0005519, DHJPAR0011537, DHJPAR0008404, DHJPAR0036528, DHJPAR0048678, 04-SRNP-14819, 07-SRNP-58539, 08-SRNP-55459, 03-SRNP-27474, 06-SRNP-30815, 05-SRNP-49341, 04-SRNP-13675, 06-SRNP-2206, 05SRNP-3370, 08-SRNP-2615, 94-SRNP-9198, 95-SRNP-11509, 95-SRNP-11286, 96-SRNP-10085, 96-SRNP10082, 96-SRNP-10080, 03-SRNP-1707, 03-SRNP-15894, 03-SRNP-10734, 96-SRNP-11563, 95-SRNP-11563, 96-SRNP-10277, 02-SRNP-5947, 02-SRNP-32717, 94-SRNP-10453, 95-SRNP-8542.1, 06-SRNP-20139. Etymology. From the Latin adjective, gracilis, meaning slender, in reference to the narrower appearance of the fly, especially of the abdomen, compared to the majority of other species. Distribution. Costa Rica, ACG, Prov. Alajuela & Guanacaste, rain forest, cloud forest, and dry forest, 17–1460 m elevation. Houghia graciloides Fleming & Wood, sp. nov. Figs. 5 b, 10 a, 24 a–f Diagnosis. Scarcely distinguishable from H. gracilis except for having sparse, minute, easily overlooked, hairs on the eyes. Description. Male. Antenna black. When viewed in profile, antenna arises distinctly above middle of eye. Length of first flagellomere extending to facial margin. Facial ridge with, in addition to the decumbent vibrissae, a row of minute, inconspicuous, hairs. Palpus pale, usually distinctly yellowish. Postgena behind postoccipital row, above level of lower facial margin, with a small patch of few black setae. Parafacial silver. Colour of fronto-orbital plate pale brassy to gold on its entire length from vertex to base of antenna (more than 50% coverage). Surface of fronto-orbital plate almost bare. Ocellar triangle, when viewed from above appearing rounded anteriorly. Diameter of anterior ocellus equal to, or greater than, diameter of base of adjacent ocellar seta. Ocellar setae arising behind anterior ocellus but closer to anterior than to posterior ocelli. Eye covered in minute hairs. Postpronotum restricted to the three main postpronotal setae. Dark stripes on either side of dorsocentral row of setae separated from one another with brownish or gray tomentosity, appearing paler than stripes, but darker than remainder of scutum. Median and lateral stripes on either side of scutum united to one another posteriorly. Postsutural dorsocentral setae 4. Anterior quadrant of anepisternum covered with short setae except for usually 3 to 5 distinctly larger setae. Katepisternum with three setae, the middle one always the smallest. Vein R1 bare dorsally. Legs ranging from reddish brown to yellow tinged but overall dark. Coxae distinctly yellow tinged, usually contrasting with darker remainder of legs. Ground colour of dorsal surface of abdominal tergite 3 and in some specimens parts of other tergites as well, dark medially but reddish laterally. Ground colour of ventral surface of abdomen partially or entirely reddish or yellowish. Sex patches present on tergite 3, as well as on tergites 4 and 5 (though sometimes small or absent on 5). Ground colour of sex patches not distinctly shiny. Terminalia: surstylus equilaterally oblong shaped, posterodorsal half haired, apex bearing many stout apical spines, tip with light inwardly apical curve when viewed dorsally. Cerci sharply pointed or distinctly tapered, apex with straight tip, ventral surface haired, separation between cerci deep broad V shape, up to 85% as long as surstylus. Lobe of sternite 5 oversized and rounded apically, inner margin covered in dense tomentosity appearing darker than surrounding cuticle, internal edge straight, single long apical seta emanating from apex. Hosts. Houghia graciloides has been reared 62 times, from 17 species in 10 genera of primarily broadleaf monocot-eating caterpillars of Hesperiinae (Hesperiidae) in ACG dry forest, cloud forest, and rain forest, out of a sample of 6000+ caterpillars. This long list of hosts overlaps with that of Houghia gracilis, but emphasizes large and long-lived monocot leaves, instead of grasses and other short-lived monocot leaves. Holotype. ♂, CNC. Type locality: Costa Rica, Area de Conservación Guanacaste, Prov. Alajuela, Sector Brasilia, Moga (11.01227°, -85.34929°), 320 m, 02/16/2010, Duvalier Briceño, DHJPAR0038672. HOUGHIA OF ACG, COSTA RICA Zootaxa 3858 (1) © 2014 Magnolia Press · 53 FIGURE 24. General morphology of Houghia graciloides sp. nov. a. Dorsal habitus. b. Frontal view of head. c. Lateral habitus. d. Posterior view of male terminalia. e. Lateral view of male terminalia. f. Ventral view of sternite 5 + segments 7+8. Paratypes. 46 ♂, 69 ♀ (CNC) Costa Rica, Prov. Guanacaste & Alajuela, ACG database codes: DHJPAR0011506, DHJPAR0008408, DHJPAR0034598, DHJPAR0008400, DHJPAR0019796, DHJPAR0008386, DHJPAR0008419, DHJPAR0008379, DHJPAR0008422, DHJPAR0008409, DHJPAR0008406, DHJPAR0008381, DHJPAR0017034, DHJPAR0008393, DHJPAR0011513, DHJPAR0019791, DHJPAR0016633, DHJPAR0011532, DHJPAR0008380, DHJPAR0008396, DHJPAR0008418, DHJPAR0008417, DHJPAR0008382, DHJPAR0038673, DHJPAR0008412, DHJPAR0008416, DHJPAR0008431, DHJPAR0008411, DHJPAR0008399, DHJPAR0008423, DHJPAR0008530, DHJPAR0008374, DHJPAR0008397, DHJPAR0008398, DHJPAR0008391, DHJPAR0008385, DHJPAR0008387, DHJPAR0019742, DHJPAR0024458, DHJPAR0016686, DHJPAR0016700, DHJPAR0008392, DHJPAR0034586, DHJPAR0034575, DHJPAR0008414, DHJPAR0008425, DHJPAR0011505, DHJPAR0008433, DHJPAR0005517, DHJPAR0005476, DHJPAR0005518, DHJPAR0016072, DHJPAR0008375, DHJPAR0008430, DHJPAR0008421, DHJPAR0020931, DHJPAR0008394, DHJPAR0008403, DHJPAR0008413, DHJPAR0008390, DHJPAR0039326, 05-SRNP-49327, 10-SRNP-65036, 06-SRNP-57990, 03-SRNP-25897, 03-SRNP-30120, 97SRNP-4152, 04-SRNP-15114, 04-SRNP-15115, 04-SRNP-14328, 04-SRNP-35037, 98-SRNP-6801, 07-SRNP31926, 07-SRNP-30976, 08-SRNP-70126, 95-SRNP-30603, 03-SRNP-30705, 07-SRNP-31935, 04-SRNP-13863, 96-SRNP-6801, 02-SRNP-4456, 02-SRNP-14318, 06-SRNP-58877, 09-SRNP-30562, 09-SRNP-30567, 05SRNP-48893, 05-SRNP-49318, 03-SRNP-1440, 03-SRNP-27125, 10-SRNP-65037, 04-SRNP-41654. Etymology. From the Latin adjective, “gracilis”, meaning slender, plus the Greek suffix, -oides, meaning resembling, in reference to its similarity to the previous species, H. gracilis. 54 · Zootaxa 3858 (1) © 2014 Magnolia Press FLEMING ET AL. Distribution. Costa Rica, ACG, Prov. Alajuela & Guanacaste, rain forest, cloud forest, and dry forest, 17–1060 m elevation. Houghia griseifrons Fleming & Wood, sp. nov. Figs. 5 c, 25 a–f Diagnosis. This small species is one of four with vein R1 setose along its entire dorsal surface, and that lack sex patches on the ventral surface of the abdomen (the other three are H. macilenta, H. nigrofemur, and H. ochrofemur). Houghia griseifrons differs from H. macilenta by the grayish rather than yellowish tomentose frons, and the lack of tomentosity on the frontal vitta leaving the ocellar triangle separated from the fronto-orbital plates by the dark brown Y-shaped upper end of the frontal vitta. FIGURE 25. General morphology of Houghia griseifrons sp. nov. a. Dorsal habitus. b. Frontal view of head. c. Lateral habitus. d. Posterior view of male terminalia. e. Lateral view of male terminalia. f. Ventral view of sternite 5 + segments 7+8. Description. Male. Antenna black. When viewed in profile, antenna arises distinctly above middle of eye. Length of first flagellomere extending to facial margin. Facial ridge bare except for a few (usually 3–5) decumbent small setae above vibrissa. Palpus pale, usually distinctly yellowish. Postgena behind postoccipital row, above level of lower facial margin, without black setae. Parafacial silver. Colour of fronto-orbital plate pale brassy to gold on its entire length from vertex to base of antenna (more than 50% coverage). Surface of fronto-orbital plate almost HOUGHIA OF ACG, COSTA RICA Zootaxa 3858 (1) © 2014 Magnolia Press · 55 bare. Ocellar triangle, when viewed from above appearing to be sharply pointed anteriorly. Diameter of anterior ocellus equal to, or greater than, diameter of base of adjacent ocellar seta. Ocellar setae arising behind anterior ocellus but closer to anterior than to posterior ocelli. Eye bare. Postpronotum restricted to the three main postpronotal setae. Dark stripes on either side of dorsocentral row of setae separated from one another with brownish or gray tomentosity, appearing paler than stripes, but darker than remainder of scutum. Median and lateral stripes on either side of scutum united to one another posteriorly. Postsutural dorsocentral setae 3, with a gap between first and third seta. Anterior quadrant of anepisternum covered with short setae except for usually 3 to 5 distinctly larger setae. Katepisternum with two setae. Vein R1 with a row of small recumbent setae along its entire length. Legs ranging from reddish brown to yellow tinged but overall dark. Coxae distinctly yellow tinged, usually contrasting with darker remainder of legs. Ground colour of dorsal surface of abdomen dark to black. Ground colour of ventral surface of abdomen entirely black. Sex patches absent. Terminalia: surstylus equilaterally oblong shaped, posterodorsal half haired, apex bearing few short apical spines, tip with light inwardly apical curve when viewed dorsally. Cerci sharply pointed or distinctly tapered, apex with straight tip, ventral surface bare, separation between cerci straight, up to 85% as long as surstylus. Lobe of sternite 5 small and pointed apically, inner margin covered in dense tomentosity appearing darker than surrounding cuticle, internal edge inwardly curved, multiple apical setae emanating from lobe. Hosts. Houghia griseifrons has been reared 3 times, from a sample of 44 dry forest and rain forest Acraga coa Schaus (Dalceridae) caterpillars. Holotype. ♂, CNC. Type locality: Costa Rica, Area de Conservación Guanacaste, Prov. Guanacaste, Sector Santa Rosa, Vado Cuajiniquil (10.94041°, -85.68043°), 275 m, 01/17/2006, Guillermo Pereira, DHJPAR0005549. Paratypes. 29 ♂, 18 ♀ (CNC) Costa Rica, Prov. Guanacaste, ACG database codes: DHJPAR0008840, 06SRNP-12341, 00-SRNP-20050. 1 ♂, USA “AZ Pima Co./ Box Canyon/ 20.IX.2001/ G. & M. Wood” and 1 ♀, USA “Congress, ARIZ./ Yavapai Co./ 23-26.IV.67/ D. M. Wood”. Etymology. From the Latin adjective, “griseus”, meaning gray and the noun, “frons”, meaning forehead, in reference to the grey tomentosity of the frons, a unique character in Houghia where the fronto-orbital plate is otherwise either silver or gold, not grey. The name is thus a noun in apposition. Distribution. Costa Rica, ACG, Prov. Guanacaste, dry forest, 275–295 m elevation, and Pima and Yavapai counties, Arizona, USA. Houghia inflatipalpis Fleming & Wood, sp. nov. Fig. 26 a–f Diagnosis. No external features have yet been found that distinguish males of H. inflatipalpis from H. marini. The surstylus of H. inflatipalpis is, however, noticably longer than the cerci, extending beyond the apex of the cerci by at least the width of the surstylus (Fig. 19e). Females of H. inflatipalpis, however, have swollen palpi (hence the name) unlike females of H. marini and most other species of Houghia. Description. Male. Antenna black. When viewed in profile, antenna arises distinctly above middle of eye. Length of first flagellomere extending to facial margin. Facial ridge bare except for a few (usually 3–5) decumbent small setae above vibrissa. Palpus pale [in females appearing swollen at its tip], usually distinctly yellowish. Postgena behind postoccipital row, above level of lower facial margin, with a small patch of few black setae. Parafacial silver. Colour of fronto-orbital plate gold only at vertex, adjacent to ocellar triangle, the remainder silver (up to 25% coverage). Surface of fronto-orbital plate almost bare. Ocellar triangle, when viewed from above appearing to be notched anteriorly. Diameter of anterior ocellus equal to, or greater than, diameter of base of adjacent ocellar seta. Ocellar setae arising behind anterior ocellus but closer to anterior than to posterior ocelli. Eye bare. Postpronotum with 4 or 5 postpronotal setae. Dark stripes on either side of dorsocentral row of setae separated from one another by yellow tomentosity. Median and lateral stripes on either side of scutum separate from each other posteriorly. Postsutural dorsocentral setae 4. Anterior quadrant of anepisternum covered with short setae except for usually 3 to 5 distinctly larger setae. Katepisternum with three setae, the middle one always the smallest. Vein R1 bare dorsally. Legs ranging from reddish brown to yellow tinged but overall dark. Coxae dark usually concolourous with remainder of leg. Ground colour of dorsal surface of abdomen dark to black. Ground colour of ventral surface of abdomen entirely black. Sex patches present on tergites 4 and 5. Ground colour of sex 56 · Zootaxa 3858 (1) © 2014 Magnolia Press FLEMING ET AL. patches shiny black. Terminalia: surstylus bayonet shaped, posterodorsal half bare, apex bearing few short apical spines, tip with light inwardly apical curve when viewed dorsally. Cerci sharply pointed or distinctly tapered, apex with blunt, hooked tip, ventral surface haired, separation between cerci straight, up to 85% as long as surstylus. Lobe of sternite 5 small and pointed apically, inner margin covered in dense tomentosity appearing darker than surrounding cuticle, internal edge inwardly curved, single long apical seta emanating from apex. FIGURE 26. General morphology of Houghia inflatipalpis sp. nov. a. Dorsal habitus. b. Frontal view of head. c. Lateral habitus. d. Posterior view of male terminalia. e. Lateral view of male terminalia. f. Ventral view of sternite 5 + segments 7+8. Hosts. Houghia inflatipalpis has been reared 56 times from 749 wild-caught caterpillars of rain forest Celaenorrhinus Burns01 and 4 times from from a sample of 266 dry forest wild-caught Celaenorrhinus eligius (Stoll) (Hesperiidae) caterpillars, all feeding on Acanthaceae; none have been reared from any other caterpillars, yet there are three other species of Celaenorrhinus Hübner, in the same two ecosystems (1041 rearings). Holotype. ♂, CNC. Type locality: Costa Rica, Area de Conservación Guanacaste, Prov. Guanacaste, Sector Pitilla, Sendero Cueston (10.99455°, -85.41461°), 640 m, 02/19/2007, Duvalier Briceño, DHJPAR0019777. Paratypes. 51 ♂, 37 ♀ (CNC) Costa Rica, Prov. Alajuela & Guanacaste, ACG database codes: DHJPAR0019667, DHJPAR0008128, DHJPAR0021020, DHJPAR0010327, DHJPAR0019720, DHJPAR0019670, DHJPAR0024495, DHJPAR0005485, DHJPAR0011488, DHJPAR0005456, DHJPAR0016617, DHJPAR0016618, DHJPAR0016606, DHJPAR0016500, DHJPAR0008504, DHJPAR0008125, DHJPAR0008494, DHJPAR0008495, DHJPAR0008497, DHJPAR0008503, DHJPAR0008501, DHJPAR0048658, DHJPAR0040682, DHJPAR0008506, DHJPAR0008499, DHJPAR0008507, DHJPAR0008498, DHJPAR0008505, DHJPAR0005560, DHJPAR0008500, HOUGHIA OF ACG, COSTA RICA Zootaxa 3858 (1) © 2014 Magnolia Press · 57 DHJPAR0008229, DHJPAR0008130, DHJPAR0011534, DHJPAR0019668, DHJPAR0037565, DHJPAR0016608, DHJPAR0016622, DHJPAR0016601, DHJPAR0029637, DHJPAR0011525, DHJPAR0019632, DHJPAR0019689, DHJPAR0040733, DHJPAR0037261, DHJPAR0037306, DHJPAR0008496, DHJPAR0008502, 06-SRNP-8517, 09-SRNP-73005, 07-SRNP-41250, 07-SRNP-1165, 08-SRNP-1253, 05-SRNP-43061, 93-SRNP-7384, 99-SRNP13361, 95-SRNP-10639, 08-SRNP-5015, 04-SRNP-60567, 04-SRNP-30777, 02-SRNP-19302, 01-SRNP-5048, 07-SRNP-2200, 99-SRNP-12313, 04-SRNP-55667, 00-SRNP-12865, 03-SRNP-10036, 02-SRNP-7747, 02SRNP-21439, 02-SRNP-19302, 06-SRNP-8521, 06-SRNP-8615, 06-SRNP-8614, 06-SRNP-8091, 07-SRNP1468, 07-SRNP-2282, 09-SRNP-33184, 07-SRNP-32393, 05-SRNP-6211. Etymology. From the Latin participle, “inflatus”, derived from the verb, “inflo” to blow into (inflate), and the noun “palpus”, literally the palm of the hand (palpus), in reference to the swollen palpi of the female of this species. Distribution. Costa Rica, ACG, Prov. Alajuela & Guanacaste, rain forest (primarily) and dry forest, 280–900 m elevation. Houghia latigena Fleming & Wood, sp. nov. Fig. 27 a–f Diagnosis. Houghia latigena is one of four nearly identical species described here, which form the H. latigena group (H. longicercus, H. latigena, H. latilobus, and H. velutina), in which the gena is higher than that of the remaining species, being 1/10 the height of the head (Figs. 27–28c, 30c & 45c) instead of 1/12 or less. In posterior view the cerci of H. latigena is long and slender, the V-shaped dorsal notch is only 1/4 or less the height of the cerci, and instead of tapering evenly (as in H. velutina) the cerci is slightly constricted in the middle (Fig. 27d). Description. Male. Antenna black. When viewed in profile, antenna arises approximately at level of middle of eye. Length of first flagellomere extending to facial margin. Facial ridge bare except for a few (usually 3–5) decumbent small setae above vibrissa. Palpus pale, usually distinctly yellowish. Postgena behind postoccipital row, above level of lower facial margin, with a small patch of few black setae. Parafacial silver. Colour of fronto-orbital plate pale brassy to gold on its entire length from vertex to base of antenna (more than 50% coverage). Surface of fronto-orbital plate covered with small recumbent hairs, especially near margin of eye. Ocellar triangle when viewed from above appearing to be notched anteriorly. Diameter of anterior ocellus equal to, or greater than, diameter of base of adjacent ocellar seta. Ocellar setae arising beside, or slightly in front of, anterior ocellus. Eye bare. Postpronotum restricted to the three main postpronotal setae. Dark stripes on either side of dorsocentral row of setae separated from one another by yellow tomentosity. Median and lateral stripes on either side of scutum separate from each other posteriorly. Postsutural dorsocentral setae 4. Anterior quadrant of anepisternum covered with short setae except for usually 3 to 5 distinctly larger setae. Katepisternum with three setae, the middle one always the smallest. Vein R1 bare dorsally. Legs ranging from reddish brown to yellow tinged but overall dark. Coxae dark usually concolourous with remainder of leg. Ground colour of dorsal surface of abdomen dark to black. Ground colour of ventral surface of abdomen entirely black. Sex patches present on tergites 4 and 5. Ground colour of sex patches not distinctly shiny. Terminalia: surstylus wedge shaped, posterodorsal half haired, apex bearing few short apical spines, tip straight. Cerci rounded, apex with straight tip, ventral surface haired, separation between cerci straight, about as long as surstylus. Lobe of sternite 5 large and rounded apically, inner margin covered in dense tomentosity appearing darker than surrounding cuticle, internal edge straight, single long apical seta emanating from apex. Hosts. Houghia latigena has been reared from Parabella macleannani (19X), Jemadia suekentonmiller (2X), Elbella scylla (Ménétriés) (1X) and Elbella merops (Bell) (2X) (Pyrginae, Hesperiidae) feeding on Myrtaceae, Salicaceae, Malpighiaceae and Melastomataceae in ACG rain forest, from a sample of 1000+ caterpillars of these three genera; while this may appear to be the host array of a “generalist” parasitoid, these caterpillars are very similar in visual appearance. Holotype. ♂, CNC. Type locality: Costa Rica, Area de Conservación Guanacaste, Prov. Alajuela, Sector San Cristobal, Potrero Argentina (10.89021°, -85.38803°), 520 m, 05/05/1999, Gloria Sihezar, DHJPAR0008442. Paratypes. 51 ♂, 35 ♀ (CNC) Costa Rica, Prov. Alajuela ACG database codes: DHJPAR0008456, DHJPAR0030175, DHJPAR0008455, DHJPAR0021985, DHJPAR0017184, DHJPAR0011510, DHJPAR0008479, 58 · Zootaxa 3858 (1) © 2014 Magnolia Press FLEMING ET AL. DHJPAR0030166, DHJPAR0008452, DHJPAR0038802, DHJPAR0039252, DHJPAR0008489, DHJPAR0008491, DHJPAR0008483, DHJPAR0008471, DHJPAR0008480, DHJPAR0008475, DHJPAR0008478, DHJPAR0008476, DHJPAR0008488, DHJPAR0008436, DHJPAR0008454, DHJPAR0008469, DHJPAR0008487, DHJPAR0008486, DHJPAR0022010, 07-SRNP-3492, 07-SRNP-32966, 09-SRNP-6934, 04-SRNP-33653, 04-SRNP-33655, 04SRNP-20837, 04-SRNP-20824, 03-SRNP-1226, 05-SRNP-21331, 05-SRNP-21363, 05-SRNP-32855, 02-SRNP14112, 03-SRNP-1829, 03-SRNP-1225, 03-SRNP-1257. Etymology. From the Latin adjective “latus”, noun, “gena”, meaning cheek, thus a noun in apposition. Distribution. Costa Rica, ACG, Prov. Alajuela & Guanacaste, rain forest, 280–560 m elevation. FIGURE 27. General morphology of Houghia latigena sp. nov. a. Dorsal habitus. b. Frontal view of head. c. Lateral habitus. d. Posterior view of male terminalia. e. Lateral view of male terminalia. f. Ventral view of sternite 5 + segments 7+8. Houghia latilobus Fleming & Wood sp. nov. Fig. 28 a–f Diagnosis. Similar to H. latigena, and part of the H. latigena group of related species in which the gena is 1/10 head height (H. longicercus, H. latigena, H. latilobus, and H. velutina). Its most distinctive feature compared to the other three species in the group is the greater width of the dorsal lobe of the cerci, and the divergent apices of the cerci (Fig. 28d). HOUGHIA OF ACG, COSTA RICA Zootaxa 3858 (1) © 2014 Magnolia Press · 59 FIGURE 28. General morphology of Houghia latilobus sp. nov. a. Dorsal habitus. b. Frontal view of head. c. Lateral habitus. d. Posterior view of male terminalia. e. Lateral view of male terminalia. f. Ventral view of sternite 5 + segments 7+8. Description. Male. Antenna black. When viewed in profile, antenna arises distinctly above middle of eye. Length of first flagellomere extending to facial margin. Facial ridge bare except for a few (usually 3–5) decumbent small setae above vibrissa. Palpus pale, usually distinctly yellowish. Postgena behind postoccipital row, above level of lower facial margin, with a small patch of few black setae. Parafacial silver. Colour of fronto-orbital plate pale brassy to gold on its entire length from vertex to base of antenna (more than 50% coverage). Surface of frontoorbital plate covered with small recumbent hairs, especially near margin of eye. Ocellar triangle, when viewed from above appearing rounded anteriorly. Diameter of anterior ocellus equal to, or greater than, diameter of base of adjacent ocellar seta. Ocellar setae arising beside, or slightly in front of, anterior ocellus. Eye bare. Postpronotum restricted to the three main postpronotal setae. Dark stripes on either side of dorsocentral row of setae separated from one another by yellow tomentosity. Median and lateral stripes on either side of scutum separate from each other posteriorly. Postsutural dorsocentral setae 4. Anterior quadrant of anepisternum covered with short setae except for usually 3 to 5 distinctly larger setae. Katepisternum with three setae, the middle one always the smallest. Vein R1 bare dorsally. Legs ranging from reddish brown to yellow tinged but overall dark. Coxae dark usually concolourous with remainder of leg. Ground colour of ventral surface of abdomen entirely black. Sex patches present on tergites 4 and 5. Ground colour of sex patches shiny black. Terminalia: surstylus wedge shaped, posterodorsal half haired, apex bearing many stout apical spines, tip with a slight outward curve. Cerci rounded, apex with blunted hook tip, ventral surface haired, separation between cerci forming a deep broad v shape, about as 60 · Zootaxa 3858 (1) © 2014 Magnolia Press FLEMING ET AL. long as surstylus. Lobe of sternite 5 large and rounded apically, inner margin covered in dense tomentosity appearing darker than surrounding cuticle, internal edge straight, single long apical seta emanating from apex. Hosts. Houghia latilobus has been reared from Elbella patrobas (Hewiston) (2X), Elbella patrobasDHJ05 (1X), Elbella scylla (Ménétriés) (1X), and Melanopyge Burns01 (1X) (Pyrginae, Hesperiidae) in ACG rain forest, from a sample of 600+ caterpillars of these two genera. Holotype. ♂, CNC. Type locality: Costa Rica, Area de Conservación Guanacaste, Prov. Guanacaste, Sector Pitilla, Medrano (11.01602°, -85.38053°), 380 m, 06/12/2012, Ricardo Calero, DHJPAR0049631. Paratypes. 2 ♂, 7 ♀ (CNC) Costa Rica, Prov. Alajuela & Guanacaste, ACG database codes: DHJPAR0016481, DHJPAR0030147, DHJPAR0021969, 08-SRNP-72327, 12-SRNP-71304, 07-SRNP-3407. Etymology. From the Latin adjective “latus”, meaning broad, and noun “lobus”, meaning hull or husk, thus a noun in apposition. Distribution. Costa Rica, ACG, Prov. Alajuela & Guanacaste, rain forest, 420–722 m elevation. Houghia longicercus Fleming & Wood, sp. nov. Fig. 29 a–f Diagnosis. Similar to H. latigena, and part of the H. latigena group of related species in which the gena is 1/10 head height (H. longicercus, H. latigena, H. latilobus, and H. velutina). Its most distinctive feature compared to the other three species in the group is the elongate cerci and relatively small dorsal V-shaped notch (Fig. 30d, and key couplets 26 and 27). Description. Male. Antenna black. When viewed in profile, antenna arises distinctly above middle of eye. Length of first flagellomere extending to facial margin. Facial ridge bare except for a few (usually 3–5) decumbent small setae above vibrissa. Palpus pale, usually distinctly yellowish. Postgena behind postoccipital row, above level of lower facial margin, with a small patch of few black setae. Parafacial silver. Colour of fronto-orbital plate pale brassy to gold on its entire length from vertex to base of antenna (more than 50% coverage). Surface of frontoorbital plate covered with small recumbent hairs, especially near margin of eye. Ocellar triangle, when viewed from above appearing to be notched anteriorly. Diameter of anterior ocellus equal to, or greater than, diameter of base of adjacent ocellar seta. Ocellar setae arising beside, or slightly in front of, anterior ocellus. Eye bare. Postpronotum restricted to the three main postpronotal setae. Dark stripes on either side of dorsocentral row of setae separated from one another by yellow tomentosity. Median and lateral stripes on either side of scutum separate from each other posteriorly. Postsutural dorsocentral setae 4. Anterior quadrant of anepisternum covered with short setae except for usually 3 to 5 distinctly larger setae. Katepisternum with three setae, the middle one always the smallest. Vein R1 bare dorsally. Legs ranging from reddish brown to yellow tinged but overall dark. Coxae dark usually concolourous with remainder of leg. Ground colour of dorsal surface of abdomen dark to black. Ground colour of ventral surface of abdomen entirely black. Sex patches present on tergites 4 and 5. Ground colour of sex patches not distinctly shiny. Terminalia: surstylus wedge shaped, posterodorsal half haired, apex bearing few short apical spines, tip straight. Cerci rounded, apex with straight tip, ventral surface haired, separation between cerci straight, about as long as surstylus. Lobe of sternite 5 large and rounded apically, inner margin covered in dense tomentosity appearing darker than surrounding cuticle, internal edge straight, single long apical seta emanating from apex. Hosts. Houghia longicercus has been reared 36 times, from a total of 917 caterpillars of Pyrrhopyge zenodorus Godman & Salvin (Pyrginae, Hesperiidae) feeding on rain forest Vismia baccifera Vandelli (Hypericaceae) and 10 times from a total of 153 Pyrrhopyge crida (Hewiston), caterpillars feeding on rain forest Vismia billbergiana Beurl. It has been reared from no other species of ACG caterpillar, but there are two other very similar species of Houghia (H. velutina and H. romeroae, see below), using other species of food plants and very similar caterpillars. Holotype. ♂, CNC. Type locality: Costa Rica, Area de Conservación Guanacaste, Prov. Guanacaste, Sector Pitilla, Loaiciga (11.01983°, -85.41342°), 445 m, 12/01/2003, Petrona Rios, DHJPAR0008473. Paratypes. 43 ♂, 43 ♀ (CNC) Costa Rica, Prov. Guanacaste & Alajuela, ACG database codes: DHJPAR0021021, DHJPAR0008450, DHJPAR0021035, DHJPAR0008462, DHJPAR0008473, DHJPAR0027897, DHJPAR0008463, DHJPAR0008481, DHJPAR0024625, DHJPAR0027858, DHJPAR0037552, DHJPAR0019589, DHJPAR0008447, DHJPAR0008484, DHJPAR0035903, DHJPAR0008492, DHJPAR0029701, DHJPAR0008465, HOUGHIA OF ACG, COSTA RICA Zootaxa 3858 (1) © 2014 Magnolia Press · 61 DHJPAR0008449, DHJPAR0008446, DHJPAR0008490, DHJPAR0008472, DHJPAR0008464, DHJPAR0008468, DHJPAR0008453, DHJPAR0008460, DHJPAR0008474, DHJPAR0008443, DHJPAR0008439, DHJPAR0024454, DHJPAR0008440, DHJPAR0008457, DHJPAR0008461, DHJPAR0008485, DHJPAR0008477, DHJPAR0008493, DHJPAR0008449, DHJPAR0008467, DHJPAR0008437, DHJPAR0008482, DHJPAR0008451, DHJPAR0008458, DHJPAR0008459, DHJPAR0008466, DHJPAR0008438, DHJPAR0008470, DHJPAR0008441, 00-SRNP-14472, 00-SRNP-20509, 94-SRNP-9325, 00-SRNP-20514, 04-SRNP-32813, 00-SRNP-14836, 04-SRNP-33914, 03SRNP-37153, 95-SRNP-494, 00-SRNP-20631, 96-SRNP-1079, 97-SRNP-6583, 07-SRNP-20895, 00-SRNP20507, 07-SRNP-32120, 03-SRNP-10574, 09-SRNP-68452. Etymology. From the Latin adjective, “latus” meaning broad, and the modern Latin noun “cercus”, meaning tail, thus a noun in apposition. Distribution. Costa Rica, ACG, Prov. Alajuela & Guanacaste, rain forest (primarily) and dry forest, 123–675 m elevation. FIGURE 29. General morphology of Houghia longicercus sp. nov. a. Dorsal habitus. b. Frontal view of head. c. Lateral habitus. d. Posterior view of male terminalia. e. Lateral view of male terminalia. f. Ventral view of sternite 5 + segments 7+8. 62 · Zootaxa 3858 (1) © 2014 Magnolia Press FLEMING ET AL. Houghia longipilosa Fleming & Wood, sp. nov. Figs. 4 b, 10 b, 30 a–f Diagnosis. One of three species with long eye hairs, it may be distinguished from H. aurifera by the silver parafacial, and from H. brevipilosa by the distinct black apical bands on tergites 2 to 5. Description. Male. Antenna black. When viewed in profile, antenna arises approximately at level of middle of eye. Length of first flagellomere shorter than facial margin (usually shorter by at least the length of the pedicel). Facial ridge bare except for a few (usually 3–5) decumbent small setae above vibrissa. Palpus pale, usually distinctly yellowish. Postgena behind postoccipital row, above level of lower facial margin, with a small patch of few black setae. Parafacial silver. Colour of fronto-orbital plate pale brassy to gold on its entire length from vertex to base of antenna (more than 50% coverage). Surface of fronto-orbital plate covered with small recumbent hairs, especially near margin of eye. Ocellar triangle, when viewed from above appearing rounded anteriorly. Diameter of anterior ocellus less than diameter of base of adjacent ocellar seta. Ocellar setae arising beside, or slightly in front of, anterior ocellus. Eye with conspicuous hairs. Postpronotum with 4 or 5 postpronotal setae. Dark stripes on either side of dorsocentral row of setae separated from one another by yellow tomentosity. Median and lateral stripes on either side of scutum separate from each other posteriorly. Postsutural dorsocentral setae 4. Anterior quadrant of anepisternum covered with long setae, none distinctly larger than the others or forming a row. Katepisternum with three setae, the middle one always the smallest. Vein R1 bare dorsally. Legs ranging from reddish brown to yellow tinged but overall dark. Coxae dark usually concolourous with remainder of leg. Ground colour of dorsal surface of abdomen dark to black. Ground colour of ventral surface of abdomen entirely black. Sex patches present on tergites 4 and 5. Ground colour of sex patches shiny black. Terminalia: surstylus wedge shaped, posterodorsal half bare, apex bearing many stout apical spines, tip with light inwardly apical curve when viewed dorsally. Cerci rounded, apex with blunt, hooked tip, ventral surface haired, separation between cerci straight, up to 85% as long as surstylus. Lobe of sternite 5 small and pointed apically, inner margin covered in dense tomentosity appearing darker than surrounding cuticle, internal edge inwardly curved, apical seta absent. Hosts. Houghia longipilosa has been reared 91 times, from a sample of 764 Memphis proserpina (Salvin) (Nymphalidae) caterpillars feeding on rain forest Mollinedia costaricensis Donn., Mollinedia viridiflora Tul. and Mollinedia pinchotiana Perkins (Monimiaceae), and one time each from the very similar caterpillars of Memphis aulica (Röber) and Memphis beatrix (Druce). Holotype. ♂, CNC. Type locality: Costa Rica, Area de Conservación Guanacaste, Prov. Guanacaste, Sector Pitilla, Sendero Laguna (10.98880°, -85.42336°), 680 m, 12/24/2004, Calixto Moraga, DHJPAR0008819. Paratypes. 118 ♂, 106 ♀ (CNC) Costa Rica, Guanacaste, ACG database codes: DHJPAR0019960, DHJPAR0046483, DHJPAR0019962, DHJPAR0008802, DHJPAR0008792, DHJPAR0008800, DHJPAR0008813, DHJPAR0008814, DHJPAR0008815, DHJPAR0008821, DHJPAR0008818, DHJPAR0008801, DHJPAR0029614, DHJPAR0008806, DHJPAR0008820, DHJPAR0008798, DHJPAR0029722, DHJPAR0008817, DHJPAR0008795, DHJPAR0008791, DHJPAR0008793, DHJPAR0008805, DHJPAR0008807, DHJPAR0008804, DHJPAR0008797, DHJPAR0008803, DHJPAR0008810, DHJPAR0008811, DHJPAR0008812, DHJPAR0011480, DHJPAR0008794, DHJPAR0008796, DHJPAR0016237, DHJPAR0029609, DHJPAR0029630, DHJPAR0008816, DHJPAR0008808, DHJPAR0016502, DHJPAR0016501, DHJPAR0017147, DHJPAR0016672, DHJPAR0011482, DHJPAR0017136, DHJPAR0016669, DHJPAR0021023, DHJPAR0040750, DHJPAR0008809, DHJPAR0029626, DHJPAR0029627, 03-SRNP-34764, 03-SRNP-34766, 03-SRNP-34392, 04-SRNP-30492, 04-SRNP-30199, 04-SRNP-30403, 04SRNP-30398, 04-SRNP-30400, 04-SRNP-30372, 04-SRNP-40250, 04-SRNP-40205, 03-SRNP-10060, 02-SRNP18339, 03-SRNP-10074, 03-SRNP-6730, 03-SRNP-6731, 02-SRNP-6024, 04-SRNP-40248, 05-SRNP-31986, 04SRNP-34297, 03-SRNP-37279, 03-SRNP-10073, 07-SRNP-36053, 06-SRNP-67825, 04-SRNP-56806, 04-SRNP56852, 06-SRNP-6043, 08-SRNP-4481, 08-SRNP-4482, 03-SRNP-37278, 06-SRNP-9936, 06-SRNP-9938, 06SRNP-9998, 99-SRNP-13648, 03-SRNP-37275, 01-SRNP-4929, 99-SRNP-12739, 99-SRNP-12328, 99-SRNP5755, 08-SRNP-4598. Etymology. From the Latin adjective, “longus”, meaning long, and the adjective “pilosus” meaning covered with hair, in reference to the hairy eyes. Although the eye hairs appear to be only slightly longer than those of H. brevipilosa or H. aurifera, they seem more conspicuous because of their greater density. Distribution. Costa Rica, ACG, Prov. Alajuela & Guanacaste, rain forest and dry forest, 320–1220 m elevation. HOUGHIA OF ACG, COSTA RICA Zootaxa 3858 (1) © 2014 Magnolia Press · 63 FIGURE 30. General morphology of Houghia longipilosa sp. nov. a. Dorsal habitus. b. Frontal view of head. c. Lateral habitus. d. Posterior view of male terminalia. e. Lateral view of male terminalia. f. Ventral view of sternite 5 + segments 7+8. Houghia luteiventris Fleming & Wood sp. nov. Fig. 31 a–f Diagnosis. This is the only species of Houghia in which the ground colour of the abdomen is translucent yellow and there are only 2 katepisternal setae. Both sexes may be recognized by these two character states. Setae on the male sex patches are black, and confined to tergites 4 and 5 (they are paler and present also on tergite 3 in H. gracilis and H. graciloides). Description. Male. Antenna black. When viewed in profile, antenna arises approximately at level of middle of eye. Length of first flagellomere almost extending to facial margin (usually shorter by less than length of pedicel). Facial ridge bare except for a few (usually 3–5) decumbent small setae above vibrissa. Palpus pale, usually distinctly yellowish. Postgena behind postoccipital row, above level of lower facial margin, with a small patch of few black setae or extensively covered with setae, and fine hairs. Parafacial silver. Colour of fronto-orbital plate pale brassy to gold on its entire length from vertex to base of antenna (more than 50% coverage). Surface of frontoorbital plate covered with small recumbent hairs, especially near margin of eye. Ocellar triangle, when viewed from above appearing to be notched anteriorly. Diameter of anterior ocellus equal to, or greater than, diameter of base of adjacent ocellar seta. Ocellar setae arising beside, or slightly in front of, anterior ocellus. Eye with 64 · Zootaxa 3858 (1) © 2014 Magnolia Press FLEMING ET AL. inconspicuous hairs. Postpronotum with 4 or 5 postpronotal setae. Dark stripes on either side of dorsocentral row of setae, separated from one another by yellow tomentosity. Median and lateral stripes on either side of scutum united to one another posteriorly. Postsutural dorsocentral setae 4. Anterior quadrant of anepisternum covered with short setae except for usually 3 to 5 distinctly larger setae. Katepisternum with two setae. Vein R1 bare dorsally. Legs ranging from reddish brown to yellow tinged but overall dark. Coxae dark usually concolourous with remainder of leg. Ground colour of dorsal surface of abdominal tergite 3, and in some specimens parts of other tergites as well, dark medially but reddish laterally. Ground colour of ventral surface of abdomen partially or entirely reddish or yellowish. Sex patches present on tergites 4 and 5. Ground colour of sex patches not distinctly shiny. Terminalia, surstylus equilaterally oblong shaped, posterodorsal half bare, apex bearing many stout apical spines, tip straight. Cerci rounded, apex with blunt, hooked tip, ventral surface haired, separation between cerci deep narrow V shape, up to 85% as long as surstylus. Lobe of sternite 5 small and pointed apically, inner margin covered in dense tomentosity appearing darker than surrounding cuticle, internal edge inwardly curved, single long apical seta emanating from apex. FIGURE 31. General morphology of Houghia luteiventris sp. nov. a. Dorsal habitus. b. Frontal view of head. c. Lateral habitus. d. Posterior view of male terminalia. e. Lateral view of male terminalia. f. Ventral view of sternite 5 + segments 7+8. Hosts. Houghia luteiventris has been reared 5 times, from 3 species of rain forest Oxytenis (Saturniidae) in a total of 2155 Oxytenis Hübner, caterpillar collections, but has been captured as an adult in other parts of rainforested Costa Rica (see localities below). Holotype. ♂, CNC. Type locality: Costa Rica, Area de Conservación Guanacaste, Prov. Guanacaste, Sector Del Oro, Canyon Rio Mena (10.99616°, -85.45562°), 560 m, 08/19/2009, Lucia Rios, DHJPAR0036505. HOUGHIA OF ACG, COSTA RICA Zootaxa 3858 (1) © 2014 Magnolia Press · 65 Paratypes. 10 ♂, 5 ♀ (CNC) Costa Rica, Prov. Guanacaste, ACG database codes: DHJPAR0008534, DHJPAR0019538, DHJPAR0027790, DHJPAR0017037, DHJPAR0021186, 06-SRNP-65811, 09-SRNP-22471, 03-SRNP-21186; Costa Rica, Guanacaste, P.N. Guanacaste, Sector Pitilla, 9 km S. Santa Cecilia, 700m, Jun 1994, C. Moraga, L N 330200_3800200 #3002; Costa Rica, Prov. Cartago, P.N. Barbilla Camino a Valle Escondido, 2 km S. Rio Dantas, 700m, 24 May 2000, E. Rojas, Manual L.N. 218200_594400 #63172; Costa Rica, Prov. Cartago, P.N. Barbilla, Turrialba, Tayutic, Campamento 2, 1200m, 5-9 May 2005, D. Briceño, M. Moraga, Libre, L_N_213371_600782 #80271; Costa Rica, Prov. Punt, Rancho Quemado, 200 m, Peninsula de Osa, 12 a 31 mar 1993. A. Gutiérrez. L-S-292500, 511000. Etymology. From the Latin adjective, “luteus”, yellow, and the feminine gender, “ventris”, of the noun “venter”, meaning belly, in reference to the pale yellow ground colour of the abdomen, particularly prominent on the underside. Distribution. Costa Rica, Puntarenas Province north to Cartago, and ACG (Prov. Alajuela & Guanacaste), rain forest, 200–1200 m elevation. Houghia macilenta Fleming & Wood, sp. nov. Fig. 32 a–f Diagnosis. This small species is one of four with vein R1 setose along its entire dorsal surface, and that lack sex patches on the ventral surfaces of the abdomen (the other three are H. griseifrons, H. nigrofemur, and H. ochrofemur). A unique feature of H. macilenta is the yellowish tomentosity along the edges of the frontal vitta, leaving only a narrow brown stripe down the middle, and entirely obscuring it between fronto-orbital plates and ocellar triangle. Description. Male. Pedicel and upper portion of first flagellomere pale orange. When viewed in profile, antenna arises distinctly above middle of eye. Length of first flagellomere extending to facial margin. Facial ridge bare except for a few (usually 3–5) decumbent small setae above vibrissa. Palpus pale, usually distinctly yellowish. Postgena behind postoccipital row, above level of lower facial margin, without black setae. Parafacial silver. Colour of fronto-orbital plate pale brassy to gold on its entire length from vertex to base of antenna (more than 50% coverage). Surface of fronto-orbital plate almost bare. Ocellar triangle, when viewed from above appearing to be sharply pointed anteriorly. Diameter of anterior ocellus equal to, or greater than, diameter of base of adjacent ocellar seta. Ocellar setae arising beside, or slightly in front of, anterior ocellus. Eye bare. Postpronotum restricted to the three main postpronotal setae. Dark stripes on either side of dorsocentral row of setae separated from one another with brownish or gray tomentosity, appearing paler than stripes, but darker than remainder of scutum. Median and lateral stripes on either side of scutum united to one another posteriorly. Postsutural dorsocentral setae 3, with a gap between first and third seta. Anterior quadrant of anepisternum covered with short setae except for usually 3 to 5 distinctly larger setae. Katepisternum with two setae. Vein R1 with a row of small recumbent setae along its entire length. Legs ranging from reddish brown to yellow tinged but overall dark. Coxae distinctly yellow tinged, usually contrasting with darker remainder of legs. Ground colour of dorsal surface of abdomen dark to black. Ground colour of ventral surface of abdomen entirely black. Sex patches absent. Terminalia: surstylus equilaterally oblong shaped, posterodorsal half haired, apex bearing few short apical spines, tip with light inwardly apical curve when viewed dorsally. Cerci sharply pointed or distinctly tapered, apex with blunt, hooked tip, ventral surface bare, separation between cerci deep narrow V shape, up to 85% as long as surstylus. Lobe of sternite 5 small and pointed apically, margins bare or not so densely pollinose as to be darkened, internal edge inwardly curved, multiple apical setae emanating from lobe. Hosts. Houghia macilenta has been reared 4 times in ACG rain forest, only from Dalceridae: Acraga hamata Schaus (2X) and two species of Dalcerides Neumogen & Dyar, among a sample of 600+ dalcerid caterpillars. Holotype. ♂, CNC. Type locality: Costa Rica, Area de Conservación Guanacaste, Prov. Guanacaste, Sector Pitilla, Pasmompa (11.01926°, -85.40997°), 440 m, 07/19/2006, Manuel Rios, DHJPAR0021031. Paratypes. 20 ♂, 12 ♀ (CNC) Costa Rica, Prov. Alajuela & Guanacaste, ACG database codes: DHJPAR0021007, DHJPAR0010482, DHJPAR0040153, 06-SRNP-33004, 07-SRNP-32683, 10-SRNP-41667, 07SRNP-42025. 66 · Zootaxa 3858 (1) © 2014 Magnolia Press FLEMING ET AL. FIGURE 32. General morphology of Houghia macilenta sp. nov. a. Dorsal habitus. b. Frontal view of head. c. Lateral habitus. d. Posterior view of male terminalia. e. Lateral view of male terminalia. f. Ventral view of sternite 5 + segments 7+8. Etymology. From the Latin adjective, “macilentus”, meaning thin, meagre, in reference to the extremely slender gena and to the narrow dark central stripe of frontal vitta bordered by golden tomentosity on the frontal vitta itself. Distribution. Costa Rica, ACG, Prov. Alajuela & Guanacaste, rain forest and dry forest, 410–440 m elevation. Houghia marini Fleming & Wood, sp. nov. Figs. 5 a, 33 a–f Diagnosis. Houghia marini is part of a group of sibling species with few distinctive features except in the male terminalia and the extent of the gold tomentosity on the fronto-orbital plate. It belongs to the H. blancoi group (H. blancoi, H. romeroae, H. chavarriae, H. marini, and H. matarritai). The eye is bare and the antenna is entirely black. The ground colour of the abdomen is black, and sex patches are confined to tergites 4 and 5. Gold tomentosity on the frons extends from vertex only to upper fronto-orbital setae (Fig. 33a) and the apex of the first flagellomere extends to the lower facial margin. This is the only member of the group in which this last character state is present. HOUGHIA OF ACG, COSTA RICA Zootaxa 3858 (1) © 2014 Magnolia Press · 67 FIGURE 33. General morphology of Houghia marini sp. nov. a. Dorsal habitus. b. Frontal view of head. c. Lateral habitus. d. Posterior view of male terminalia. e. Lateral view of male terminalia. f. Ventral view of sternite 5 + segments 7+8. Description. Male. Antenna black. When viewed in profile, antenna arises distinctly above middle of eye. Length of first flagellomere extending to facial margin. Facial ridge bare except for a few (usually 3–5) decumbent small setae above vibrissa. Palpus pale, usually distinctly yellowish. Postgena behind postoccipital row, above level of lower facial margin, with a small patch of few black setae. Parafacial silver. Colour of fronto-orbital plate gold only at vertex, adjacent to ocellar triangle, the remainder silver (up to 25% coverage). Surface of fronto-orbital plate almost bare. Ocellar triangle when viewed from above appearing to be notched anteriorly. Diameter of anterior ocellus equal to, or greater than, diameter of base of adjacent ocellar seta. Ocellar setae arising behind anterior ocellus but closer to anterior than to posterior ocelli. Eye bare. Postpronotum with 4 or 5 postpronotal setae. Dark stripes on either side of dorsocentral row of setae separated from one another with brownish or gray tomentosity, appearing paler than stripes, but darker than remainder of scutum. Median and lateral stripes on either side of scutum united to one another posteriorly. Postsutural dorsocentral setae 4. Anterior quadrant of anepisternum covered with short setae except for usually 3 to 5 distinctly larger setae. Katepisternum with three setae, the middle one always the smallest. Vein R1 bare dorsally. Legs ranging from reddish brown to yellow tinged but overall dark. Coxae dark usually concolourous with remainder of leg. Ground colour of dorsal surface of abdomen dark to black. Ground colour of ventral surface of abdomen entirely black. Sex patches present on tergites 4 and 5. Ground colour of sex patches shiny black. Terminalia: surstylus equilaterally oblong shaped, posterodorsal 68 · Zootaxa 3858 (1) © 2014 Magnolia Press FLEMING ET AL. half bare, apex bearing many stout apical spines, tip with light inwardly apical curve when viewed dorsally. Cerci rounded, apex with blunt, hooked tip, ventral surface haired, separation between cerci deep broad V shape, about as long as surstylus. Lobe of sternite 5 small and pointed apically, inner margin covered in dense tomentosity appearing darker than surrounding cuticle, internal edge inwardly curved, multiple apical setae emanating from lobe. Hosts. Houghia marini has been reared 67 times, from wild-caught rain forest caterpillars of 3 species of Euselasia Hübner (Riodinidae). More than 2170 of the caterpillars of these Euselasia species have been collected; however, since they are gregarious feeders as caterpillars, we note that H. marini was reared from 11 of 41 groups of caterpillars. All H. marini rearings are from rain forest, even though the host caterpillars occur in both rain forest and dry forest. Holotype. ♂, CNC. Type locality: Costa Rica, Area de Conservación Guanacaste, Prov. Alajuela, Sector San Cristobal, Sendero Colegio (10.89296°, -85.37880°), 520 m, 04/09/2008, Elda Araya, DHJPAR0024511. Paratypes. 25 ♂, 29 ♀ (CNC) Costa Rica, Prov. Guanacaste, ACG database codes: DHJPAR0042658, DHJPAR0042287, DHJPAR0042281, DHJPAR0042282, DHJPAR0042283, DHJPAR0042284, DHJPAR0008157, DHJPAR0008131, DHJPAR0008122, DHJPAR0008116, DHJPAR0008158, DHJPAR0008120, DHJPAR0008124, DHJPAR0035676, DHJPAR0035682, DHJPAR0035901, DHJPAR0035905, DHJPAR0010046, DHJPAR0024500, DHJPAR0024501, DHJPAR0021862, DHJPAR0021859, DHJPAR0021860, DHJPAR0008145, DHJPAR0008147, DHJPAR0008149, DHJPAR0008150, DHJPAR0008152, DHJPAR0008153, DHJPAR0008154, DHJPAR0008155, DHJPAR0008156, DHJPAR0008144, DHJPAR0008556, DHJPAR0008564, DHJPAR0008560, DHJPAR0008562, DHJPAR0008159, DHJPAR0008561, DHJPAR0007152, DHJPAR0007160, DHJPAR0008558, DHJPAR0008559, DHJPAR0042285, DHJPAR0008563, DHJPAR0035910, DHJPAR0021858, DHJPAR0042286, DHJPAR0024502, DHJPAR0024507, DHJPAR0024504, DHJPAR0008146, DHJPAR0021861, DHJPAR0024506, DHJPAR0008151, DHJPAR0024512, 99-SRNP-5208, 02-SRNP-33405. Etymology. Houghia marini is dedicated to Sigifredo Marin of Liberia, Guanacaste, Costa Rica, in recognition of his decades of enthusiastic administration of ACG and of GDFCF field projects in ACG, and special attention to the parataxonomists rearing the tachinids described here. Distribution. Costa Rica, ACG, Prov. Guanacaste, rain forest, 280–520 m elevation. Houghia matarritai Fleming & Wood, sp. nov. Fig. 34 a–f Diagnosis. Houghia matarritai belongs to the H. blancoi group (H. blancoi, H. romeroae, H. chavarriae, H. marini, and H. matarritai). It has few distinctive features except in the male terminalia and the presence of gold tomentosity covering in excess of 50% of the fronto-orbital plate. The eye is bare and the antennae are entirely black. The ground colour of the abdomen is black, and sex patches are confined to tergites 4 and 5. Gold tomentosity extends from the vertex to lower fronto-orbital setae, but the apex of the first flagellomere falls short of the lower facial margin by half the length of the pedicel (Fig. 34b). Description. Male. Antenna black. When viewed in profile, antenna arises distinctly above middle of eye. Length of first flagellomere almost extending to facial margin, (usually shorter by less than length of pedicel). Facial ridge bare except for a few (usually 3–5) decumbent small setae above vibrissa. Palpus pale, usually distinctly yellowish. Postgena behind postoccipital row, above level of lower facial margin, with a small patch of few black setae. Parafacial silver. Colour of fronto-orbital plate pale brassy to gold on its entire length from vertex to base of antenna (more than 50% coverage). Surface of fronto-orbital plate covered with small recumbent hairs, especially near margin of eye. Ocellar triangle appearing rounded anteriorly when viewed from above. Diameter of anterior ocellus less than diameter of base of adjacent ocellar seta. Ocellar setae arising beside, or slightly in front of, anterior ocellus. Eye with inconspicuous hairs. Postpronotum with 4 or 5 postpronotal setae. Dark stripes on either side of dorsocentral row of setae separated from one another by yellow tomentosity. Median and lateral stripes on either side of scutum separate from each other posteriorly. Postsutural dorsocentral setae 4. Anterior quadrant of anepisternum covered with short setae except for usually 3 to 5 distinctly larger setae. Katepisternum with three setae, the middle one always the smallest. Vein R1 bare dorsally. Legs ranging from reddish brown to yellow tinged but overall dark. Coxae dark usually concolourous with remainder of leg. Ground colour of dorsal HOUGHIA OF ACG, COSTA RICA Zootaxa 3858 (1) © 2014 Magnolia Press · 69 surface of abdomen dark to black. Ground colour of ventral surface of abdomen entirely black. Sex patches present on tergites 4 and 5. Ground colour of sex patches shiny black. Terminalia: surstylus bayonet shaped, posterodorsal half bare, apex bearing many stout apical spines, tip with light inwardly apical curve when viewed dorsally. Cerci rounded, apex with blunt, hooked tip, ventral surface haired, separation between cerci deep narrow V shape, up to 85% as long as surstylus. Lobe of sternite 5 small and pointed apically, inner margin covered in dense tomentosity appearing darker than surrounding cuticle, internal edge inwardly curved, single long apical seta emanating from apex. FIGURE 34. General morphology of Houghia matarritai sp. nov. a. Dorsal habitus. b. Frontal view of head. c. Lateral habitus. d. Posterior view of male terminalia. e. Lateral view of male terminalia. f. Ventral view of sternite 5 + segments 7+8. Hosts. Houghia matarritai has been reared 60 times, from caterpillars of 4 species of monocot-eating Nymphalidae (Pierella Westwood, Taygetis Hübner) and 14 species of monocot-eating hesperiine Hesperiidae in ACG dry forest and rain forest, out of a sample of 4000+ caterpillars. There is one exception: the single rearing from Dunama jessiebarronae Chacón (Notodontidae) was from a monocot-eating caterpillar. It has been reared only from this ecological grouping of ACG caterpillars. Holotype. ♂, CNC. Type locality: Costa Rica, Area de Conservación Guanacaste, Prov. Alajuela, Sector Rincon Rain Forest, Sendero Venado (10.89678°, -85.27001°), 420 m, 12/08/2008, Jose Perez, DHJPAR0030019. Paratypes. 70 ♂, 74 ♀ (CNC) Costa Rica, Prov. Guanacaste, ACG database codes: DHJPAR0008557, DHJPAR0011527, DHJPAR0029799, DHJPAR0019631, DHJPAR0016604, DHJPAR0008527, DHJPAR0014998, DHJPAR0016344, DHJPAR0011526, DHJPAR0011514, DHJPAR0011518, DHJPAR0011523, DHJPAR0016614, 70 · Zootaxa 3858 (1) © 2014 Magnolia Press FLEMING ET AL. DHJPAR0008169, DHJPAR0008126, DHJPAR0008161, DHJPAR0042705, DHJPAR0008509, DHJPAR0006596, DHJPAR0011517, DHJPAR0048515, DHJPAR0011521, DHJPAR0008166, DHJPAR0008160, DHJPAR0008165, DHJPAR0008508, DHJPAR0008551, DHJPAR0008171, DHJPAR0030021, DHJPAR0008172, DHJPAR0008532, DHJPAR0027860, DHJPAR0008531, DHJPAR0011473, DHJPAR0006947, DHJPAR0014982, DHJPAR0010224, DHJPAR0017146, DHJPAR0011507, DHJPAR0007121, DHJPAR0024543, DHJPAR0011524, DHJPAR0008118, DHJPAR0011535, DHJPAR0016111, DHJPAR0008528, DHJPAR0011522, DHJPAR0011519, DHJPAR0016492, DHJPAR0008167, DHJPAR0008119, DHJPAR0008164, DHJPAR0048515, DHJPAR0011521, DHJPAR0011520, DHJPAR0008163, DHJPAR0008173, DHJPAR0008524, DHJPAR0008525, DHJPAR0010011, DHJPAR0008170, DHJPAR0008535, DHJPAR0011481, DHJPAR0008111, DHJPAR0011516, DHJPAR0008168, 05-SRNP-42616, 05-SRNP-42567, 05-SRNP-42576, 05-SRNP-42138, 05-SRNP-42142, 04-SRNP-40623, 06-SRNP-41954, 04SRNP-41183, 06-SRNP-32172, 05-SRNP-42140, 05-SRNP-42460, 04-SRNP-60960, 05-SRNP-42575, 05-SRNP42424, 05-SRNP-42427, 05-SRNP-42461, 04-SRNP-14343, 03-SRNP-12350.1, 04-SRNP-40001, 04-SRNP60760, 05-SRNP-42138, 02-SRNP-42682, 04-SRNP-60438, 01-SRNP-17731, 05-SRNP-42141, 05-SRNP-42579, 04-SRNP-40624, 07-SRNP-41428, 06-SRNP-90, 06-SRNP-43579, 06-SRNP-41954, 06-SRNP-47507, 07-SRNP66183, 05-SRNP-42425. Etymology. Houghia matarritai is dedicated to Federico Matarrita of Liberia in recognition of his development of the ACG web site (http://www.acguanacaste.ac.cr) as an encyclopedia of ACG history and activities, including the inventory of these tachinids. Distribution. Costa Rica, ACG, Prov. Guanacaste, dry forest and rain forest, 17–590 m elevation. Houghia nigrofemur Fleming & Wood, sp. nov. Fig. 9 b, 35 a–c Diagnosis. This species is one of four with vein R1 setose along its entire dorsal surface, and that lack sex patches on the ventral side of the abdomen (the other three are Houghia macilenta, H. griseifrons, and H. ochrofemur). The pedicel, and most of the first flagellomere are orange, and the latter extends well below the lower facial margin; the facial ridge is strontly curved, nearly obliterating the parafacial ventrally, as in H. ochrofemur. However, the legs are entirely black, and the first flagellomere is only slightly curved (n=1) not S-shaped as in H. ochrofemur. Description. Male. Pedicel and upper portion of first flagellomere pale orange. When viewed in profile, antenna arises distinctly above middle of eye. Length of first flagellomere extending to facial margin. First flagellomere merely slightly bowed anteriorly (only one male available). Facial ridge bare except for a few (usually 3–5) decumbent small setae above vibrissa. Palpus pale, usually distinctly yellowish. Postgena behind postoccipital row, above level of lower facial margin, without black setae. Parafacial silver. Colour of fronto-orbital plate pale brassy to gold on its entire length from vertex to base of antenna (more than 50% coverage). Surface of frontoorbital plate almost bare. Ocellar triangle, when viewed from above appearing to be sharply pointed anteriorly. Diameter of anterior ocellus equal to, or greater than, diameter of base of adjacent ocellar seta. Ocellar setae arising behind anterior ocellus but closer to anterior than to posterior ocelli. Eye bare. Postpronotum restricted to the three main postpronotal setae. Dark stripes on either side of dorsocentral row of setae separated from one another by yellow tomentosity. Median and lateral stripes on either side of scutum united to one another posteriorly. Postsutural dorsocentral setae 4. Anterior quadrant of anepisternum covered with short setae except for usually 3 to 5 distinctly larger setae. Katepisternum with two setae. Vein R1 with a row of small recumbent setae along its entire length. Legs ranging from reddish brown to yellow tinged but overall dark. Coxae dark usually concolourous with remainder of leg. Ground colour of dorsal surface of abdomen dark to black. Ground colour of ventral surface of abdomen entirely black. Sex patches absent.Terminalia for this species were not dissected as it is a readily distinguishable species and there was only one male available, which is the holotype. Dissections will be carried out at a later date when there are more specimens. Host: Houghia nigrofemur has been reared only 3 times, from Antaeotricha Zeller and Antaeocerconota (Elachistidae) feeding on 2 species of rain forest Inga Miller (Fabaceae), in a sample of 7800+ caterpillars of these two genera of small moths. Holotype. ♂, CNC. Type locality: Costa Rica, Area de Conservación Guanacaste, Prov. Guanacaste, Sector Del Oro, Margarita (11.03234°, -85.43954°), 380 m, 12/07/2007, Roster Moraga, DHJPAR0024570. HOUGHIA OF ACG, COSTA RICA Zootaxa 3858 (1) © 2014 Magnolia Press · 71 Paratypes. 2 ♀ (CNC) Costa Rica, Guanacaste, ACG database codes: DHJPAR0023659, DHJPAR0046702. Etymology. From the Latin adjective “niger”, meaning black, and the noun “femur”, meaning thigh, thus a noun in apposition. Distribution. Costa Rica, ACG, Prov. Guanacaste, rain forest, 95–380 m elevation. FIGURE 35. General morphology of Houghia nigrofemur sp. nov. a. Dorsal habitus. b. Frontal view of head. c. Lateral habitus. Houghia ochrofemur Fleming & Wood, sp. nov. Fig. 36 a–f Diagnosis. This species is one of four with vein R1 setose along its entire dorsal surface, and that lack sex patches on the ventral side of the abdomen (the other three are Houghia macilenta, H. griseifrons, and H. nigrofemur). The pedicel and most of the first flagellomere are orange and the latter is medially incurved and extends well below the lower facial margin. The facial ridge is strongly curved, nearly obliterating the parafacial ventrally. The femora are orange basally but black apically (all black in H. nigrofemur), and the tibiae and tarsi are black. Description. Male. Pedicel and upper portion of first flagellomere pale orange. When viewed in profile, antenna arises distinctly above middle of eye. Length of first flagellomere extending to facial margin. First flagellomere distinctly S-shaped, the lower curve parallelling strongly curved facial ridge. Facial ridge bare except for a few (usually 3–5) decumbent small setae above vibrissa. Palpus pale, usually distinctly yellowish. Postgena behind postoccipital row, above level of lower facial margin, without black setae. Parafacial silver. Colour of fronto-orbital plate pale brassy to gold on its entire length from vertex to base of antenna (more than 50% coverage). Surface of fronto-orbital plate almost bare. Ocellar triangle when viewed from above appearing to be sharply pointed anteriorly. Diameter of anterior ocellus equal to, or greater than, diameter of base of adjacent ocellar seta. Ocellar setae arising distinctly behind anterior ocellus, approximately midway between anterior and posterior ocelli. Eye bare. Postpronotum restricted to the three main postpronotal setae. Dark stripes on either side of dorsocentral row of setae separated from one another by yellow tomentosity. Median and lateral stripes on either side of scutum separate from each other posteriorly. Postsutural dorsocentral setae 4. Anterior quadrant of anepisternum covered with short setae except for usually 3 to 5 distinctly larger setae. Katepisternum with two setae. Vein R1 with a row of small recumbent setae along its entire length. Legs ranging from reddish brown to yellow tinged but overall dark or distinctly pale, often light yellow. Coxae distinctly yellow tinged, usually contrasting with darker remainder of legs. Ground colour of dorsal surface of abdomen dark to black. Ground colour of ventral surface of abdomen entirely black. Sex patches absent. Terminalia: surstylus bayonet shaped, 72 · Zootaxa 3858 (1) © 2014 Magnolia Press FLEMING ET AL. posterodorsal half haired, apex bearing many stout apical spines, tip with light inwardly apical curve when viewed dorsally. Cerci sharply pointed or distinctly tapered, apex with straight tip, ventral surface bare, separation between cerci deep narrow V shape, up to 85% as long as surstylus. Lobe of sternite 5 small and pointed apically, margins bare or not so densely pollinose as to be darkened, internal edge straight, multiple apical setae emanating from lobe. Hosts. Houghia ochrofemur has been reared 13 times, exclusively from 7 species of rain forest Antaeotricha Zeller (Elachistidae) feeding on 8 species of plants in 6 plant families, among a sample of 8000+ caterpillars of this genus. Holotype. ♂, CNC. Type locality: Costa Rica, Area de Conservación Guanacaste, Prov. Guanacaste, Sector Pitilla, Estacion Quica (10.99697°, -85.39666°), 470 m, 6/14/2008, Oscar Siezar, DHJPAR0027919. Paratypes. 5 ♂, 6 ♀ (CNC) Costa Rica, Prov. Alajuela and Guanacaste, ACG database codes: DHJPAR0035678, DHJPAR0049551, DHJPAR0046695, DHJPAR0035692, DHJPAR0037568, DHJPAR0036585, DHJPAR0046696, DHJPAR0036578, DHJPAR0048657, DHJPAR0036572, DHJPAR0042640. Etymology. From the Latin adjective “ochra”, meaning pale yellow, and “femur”, meaning thigh, thus a noun in apposition. Distribution. Costa Rica, ACG, Prov. Alajuela & Guanacaste, rain forest, 123–475 m elevation. FIGURE 36. General morphology of Houghia ochrofemur sp. nov. a. Dorsal habitus. b. Frontal view of head. c. Lateral habitus. d. Posterior view of male terminalia. e. Lateral view of male terminalia. f. Ventral view of sternite 5 + segments 7+8. HOUGHIA OF ACG, COSTA RICA Zootaxa 3858 (1) © 2014 Magnolia Press · 73 Houghia omissa Fleming & Wood, sp. nov. Figs. 7 b, 10 c, 37 a–f Diagnosis. One of a group of seven closely related species in ACG, and many more in Central and South America, the H. crypta group, in which the second postsutural dorsocentral seta is missing, leaving a gap between first and third setae (H. omissa, H. confinis, H. triangularis, H. destituta, H. crypta, H. biseriata, H. bivittata). It is the only species of Houghia known to us in which tergite 4 has a well developed sex patch while tergite 5 lacks any trace of a patch (Fig. 10c). FIGURE 37. General morphology of Houghia omissa sp. nov. a. Dorsal habitus. b. Frontal view of head. c. Lateral habitus. d. Posterior view of male terminalia. e. Lateral view of male terminalia. f. Ventral view of sternite 5 + segments 7+8. Description. Male. Antenna black. When viewed in profile, antenna arises approximately at level of middle of eye. Length of first flagellomere extending to facial margin. Facial ridge bare except for a few (usually 3–5) decumbent small setae above vibrissa. Palpus pale, usually distinctly yellowish. Postgena behind postoccipital row, above level of lower facial margin, with a small patch of few black setae. Parafacial silver. Surface of fronto-orbital plate almost bare. Ocellar triangle when viewed from above appearing rounded anteriorly. Diameter of anterior ocellus less than diameter of base of adjacent ocellar seta. Ocellar setae arising behind anterior ocellus but closer to anterior than to posterior ocelli. Eye bare. Postpronotum restricted to the three main postpronotal setae. Dark stripes on either side of dorsocentral row of setae separated from one another with brownish or gray tomentosity, appearing paler than stripes, but darker than remainder of scutum. Median and lateral stripes on either side of 74 · Zootaxa 3858 (1) © 2014 Magnolia Press FLEMING ET AL. scutum united to one another posteriorly. Postsutural dorsocentral setae 3, with a gap between first and third seta. Anterior quadrant of anepisternum covered with short setae except for usually 3 to 5 distinctly larger setae. Katepisternum with two setae. Vein R1 bare dorsally. Legs ranging from reddish brown to yellow tinged but overall dark. Coxae dark usually concolourous with remainder of leg. Ground colour of dorsal surface of abdomen dark to black. Ground colour of ventral surface of abdomen entirely black. Sex patches present only on tergite 4. Ground colour of sex patches shiny black. Terminalia: surstylus equilaterally oblong shaped, posterodorsal half bare, apex bearing few short apical spines, tip with light inwardly apical curve when viewed dorsally. Cerci sharply pointed or distinctly tapered, apex with blunt, hooked tip, ventral surface bare, separation between cerci straight, up to 85% as long as surstylus. Lobe of sternite 5 large and rounded apically, inner margin covered in dense tomentosity appearing darker than surrounding cuticle, internal edge straight, multiple apical setae emanating from lobe. Hosts. Houghia omissa has been reared only 6 times, and always from a sample of 47 Thracides nanea nida Evans (Hesperiinae, Hesperiidae) caterpillars feeding on the foliage of rain forest understory Asplundia microphylla (Oerst.) (Cyclanthaceae). Holotype. ♂, CNC. Type locality: Costa Rica, Area de Conservación Guanacaste, Prov. Guanacaste, Sector Pitilla, Ingas (11.00311°, -85.42041°), 580 m, 01/23/2004, Petrona Rios, DHJPAR0018946. Paratypes. 10 ♂, 16 ♀ (CNC) Costa Rica, Prov. Alajuela and Guanacaste, ACG database codes: DHJPAR0008782, DHJPAR0017125, DHJPAR0005451, DHJPAR0016603, DHJPAR0030434, 05-SRNP-6520, 98-SRNP-6786, 06-SRNP-8227, 09-SRNP-40189. Etymology. From the Latin participle of the verb “omitto”, meaning to leave out or omit, referring to the lack of a sex patch on tergite 5, although a well developed sex patch is present on tergite 4, a unique character state that defines the species. Distribution. Costa Rica, ACG, Prov. Alajuela & Guanacaste, rain forest, 420–645 m elevation. Houghia pallida Fleming & Wood, sp. nov. Fig. 38 a–f Diagnosis. A distinctive species with ground colour of abdominal tergites 1+2 and 3 translucent yellow to pale orange except for a black mid-dorsal stripe; tergites 1 to 3 with little pale tomentosity dorsally, but entirely yellow ventrally; tergites 4 and 5 mostly black dorsally with silver tomentosity laterally, but pale yellow below; hair on ventral surface of tergite 3 sparse, and could be mistaken for a sex patch (as is present in H. gracilis and H. graciloides) but hairs sparser and more erect than on tergite 4; pedicel and most of first flagellomere orange; scutal stripes on each side of scutum enclosing brown tomentosity, thus appearing as a single broad stripe (as in H. bivittata and H. biseriata). Description. Male. Pedicel and most of first flagellomere pale orange. When viewed in profile, antenna arises distinctly above middle of eye. Length of first flagellomere extending to facial margin. Facial ridge bare except for a few (usually 3–5) decumbent small setae above vibrissa. Palpus pale, usually distinctly yellowish. Postgena behind postoccipital row, above level of lower facial margin, without black setae. Parafacial silver. Colour of fronto-orbital plate gold on upper half, from vertex to midway down face (up to 50% coverage). Surface of frontoorbital plate almost bare. Ocellar triangle, when viewed from above appearing to be sharply pointed anteriorly. Diameter of anterior ocellus equal to, or greater than, diameter of base of adjacent ocellar seta. Ocellar setae arising distinctly behind anterior ocellus, approximately midway between anterior and posterior ocelli. Eye bare. Postpronotum restricted to the three main postpronotal setae. Dark stripes on either side of dorsocentral row of setae, separated from one another with brownish or gray tomentosity, appearing paler than stripes, but darker than remainder of scutum. Median and lateral stripes on either side of scutum separate from each other posteriorly. Postsutural dorsocentral setae 3, with a gap between first and third seta, middle seta usually reduced in size. Anterior quadrant of anepisternum covered with short setae except for usually 3 to 5 distinctly larger setae. Katepisternum bearing three setae, the middle one always the smallest. Vein R1 bare dorsally. Legs ranging from reddish brown to yellow tinged but overall dark. Coxae distinctly yellow tinged, usually contrasting with darker remainder of legs. Ground colour of dorsal surface of abdominal tergite 3, and in some specimens parts of other tergites as well, dark medially but reddish laterally. Ground colour of ventral surface of abdomen partially or entirely reddish or yellowish. Sex patches present on tergite 3, as well as on tergites 4 and 5 (though sometimes HOUGHIA OF ACG, COSTA RICA Zootaxa 3858 (1) © 2014 Magnolia Press · 75 small or absent on 5). Ground colour of sex patches not distinctly shiny. Terminalia: surstylus equilaterally oblong shaped, posterodorsal half haired, apex devoid of apical spines, tip straight. Cerci blunted apically in dorsal view (boxlike), apex with straight tip, ventral surface bare, separation between cerci straight, about as long as surstylus. Lobe of sternite 5 oversized and rounded apically, inner margin covered in dense tomentosity appearing darker than surrounding cuticle, internal edge straight, multiple apical setae emanating from lobe. FIGURE 38. General morphology of Houghia pallida sp. nov. a. Dorsal habitus. b. Frontal view of head. c. Lateral habitus. d. Posterior view of male terminalia. e. Lateral view of male terminalia. f. Ventral view of sternite 5 + segments 7+8. Hosts. Houghia pallida has been reared 22 times in ACG rain forest from a sample of 219 caterpillars of Aroma aroma (Hewiston) (10X) and A. henricus (Staudinger) (12X) (Hesperiinae, Hesperiidae) feeding on leaves of Cyclanthus bipartatus Poiteau (Cyclanthaceae), a perennial large herb in the shaded understory. Holotype. ♂, CNC. Type locality: Costa Rica, Area de Conservación Guanacaste, Prov. Alajuela, Sector San Cristobal, Sendero Colegio (10.89296°, -85.37880°), 520 m, 02/12/2005, Carolina Cano, DHJPAR0008432. Paratypes. 35 ♂, 27 ♀ (CNC) Costa Rica, Guanacaste, ACG database codes: DHJPAR0008520, DHJPAR0024600, DHJPAR0019529, DHJPAR0023636, DHJPAR0034502, DHJPAR0008523, DHJPAR0008373, DHJPAR0008522, DHJPAR0016151, DHJPAR0008514, DHJPAR0008510, DHJPAR0021009, DHJPAR0008511, DHJPAR0017164, DHJPAR0008518, DHJPAR0007128, DHJPAR0017044, DHJPAR0008521, DHJPAR0008515, DHJPAR0008519, DHJPAR0040659, 08-SRNP-30621, 08-SRNP-20624, 07-SRNP-32167, 09-SRNP-65479, 05SRNP-41817, 02-SRNP-19383, 05-SRNP-20768, 04-SRNP-60636, 04-SRNP-30821, 05-SRNP-65825, 07-SRNP41525, 04-SRNP-4593, 07-SRNP-40540, 02-SRNP-21562, 04-SRNP-31064, 10-SRNP-32015. 76 · Zootaxa 3858 (1) © 2014 Magnolia Press FLEMING ET AL. Etymology. From Latin, referring to the orange antenna and extensively pale abdomen, in which tergites 1+2, most of tergite 3, anterolateral corners of tergite 4, and entire ventral surface are pale orange, a pattern not yet known in other members of the genus. Distribution. Costa Rica, ACG, Prov. Guanacaste, rain forest, 320–740 m elevation. Houghia parmata Fleming & Wood, sp. nov. Fig. 39 a–f Diagnosis. The most distinctive feature of the male of H. parmata is the broad, bright silvery frons (Fig. 39b); the frontal vitta is extremely narrow and the lower reclinate orbital setae arise farther back than those of other species, nearer the posterior proclinate fronto-orbital seta, creating a broad silvery frons relatively uninterrupted by setae. FIGURE 39. General morphology of Houghia parmata sp. nov. a. Dorsal habitus. b. Frontal view of head. c. Lateral habitus. d. Posterior view of male terminalia. e. Lateral view of male terminalia. f. Ventral view of sternite 5 + segments 7+8. Description. Male. Antenna black. When viewed in profile, antenna arises distinctly above middle of eye. Length of first flagellomere almost extending to facial margin (usually shorter by less than length of pedicel). Facial ridge bare except for a few (usually 3–5) decumbent small setae above vibrissa. Palpus pale, usually distinctly yellowish. Postgena behind postoccipital row, above level of lower facial margin, with a small patch of few black setae. Parafacial silver. Colour of fronto-orbital plate pale brassy to gold on its entire length from vertex HOUGHIA OF ACG, COSTA RICA Zootaxa 3858 (1) © 2014 Magnolia Press · 77 to base of antenna (more than 50% coverage). Surface of fronto-orbital plate covered with small recumbent hairs, especially near margin of eye. Ocellar triangle, when viewed from above appearing to be notched anteriorly. Diameter of anterior ocellus less than diameter of base of adjacent ocellar seta. Ocellar setae arising beside, or slightly in front of, anterior ocellus. Eye bare. Postpronotum with 4 or 5 postpronotal setae. Dark stripes on either side of dorsocentral row of setae, separated from one another by yellow tomentosity. Median and lateral stripes on either side of scutum united to one another posteriorly. Postsutural dorsocentral setae 4. Anterior quadrant of anepisternum covered with short setae except for usually 3 to 5 distinctly larger setae. Katepisternum bearing three setae, the middle one always the smallest. Vein R1 bare dorsally. Legs ranging from reddish brown to yellow tinged but overall dark. Coxae dark, usually concolourous with remainder of leg. Ground colour of dorsal surface of abdomen dark to black. Ground colour of ventral surface of abdomen entirely black. Sex patches present on tergites 4 and 5. Ground colour of sex patches shiny black. Terminalia: surstylus equilaterally oblong shaped, posterodorsal half bare, apex bearing many stout apical spines, tip with light inwardly apical curve when viewed dorsally. Cerci rounded, apex with straight tip, ventral surface haired, separation between cerci deep broad V shape, up to 85% as long as surstylus. Lobe of sternite 5 small and pointed apically, inner margin covered in dense tomentosity appearing darker than surrounding cuticle, internal edge inwardly curved, apical seta absent. Hosts. Houghia parmata has been reared only 6 times, and always from Anomis luridula Guenée (Noctuidae) feeding on Hampea appendiculata Donnel-Smith (Malvaceae), from a sample of 487 caterpillars. Holotype. ♂, CNC. Type locality: Costa Rica, Area de Conservación Guanacaste, Prov. Guanacaste, Sector Pitilla, Leonel (10.99637°, -85.40195°), 510 m, 07/04/2008, Mauricio Siezar, DHJPAR0027895. Paratypes. 2 ♂, 5 ♀ (CNC) Costa Rica, Prov. Alajuela and Guanacaste, ACG database codes: DHJPAR0008526, DHJPAR0017131, DHJPAR0027872, DHJPAR0008533, DHJPAR0020992, DHJPAR0040667, DHJPAR0011503. Etymology. From the Latin adjective “parmata”, meaning “armed with the parma”, a small round shield, referring to the broad shiny silver frons with central area uninterrupted by setae or wide frontal vitta, that suggests a shield carried in front of the head. Distribution. Costa Rica, ACG, Prov. Alajuela & Guanacaste, rain forest, 410–525 m elevation. Houghia pilosifrons Fleming & Wood, sp. nov. Figs. 8 a, 40 a–f Diagnosis. This species lacks distinctive character states, except for the more pilose, entirely gold tomentose, fronto-orbital plate. The antenna arises at the level of the middle of head height. Ground colour of the abdomen may be either entirely black or somewhat reddish laterally at base. Apex of the surstylus in posterior view abruptly narrowed and bent medially (Fig. 40d). Description. Male. Antenna black. When viewed in profile, antenna arises approximately at level of middle of eye. Length of first flagellomere shorter than facial margin (usually shorter by at least the length of the pedicel). Facial ridge bare except for a few (usually 3–5) decumbent small setae above vibrissa. Palpus pale, usually distinctly yellowish. Postgena behind postoccipital row, above level of lower facial margin, with a small patch of few black setae. Parafacial silver. Colour of fronto-orbital plate pale brassy to gold on its entire length from vertex to base of antenna (more than 50% coverage). Surface of fronto-orbital plate covered with small recumbent hairs, especially near margin of eye. Ocellar triangle, when viewed from above appearing to be notched anteriorly. Diameter of anterior ocellus equal to, or greater than, diameter of base of adjacent ocellar seta. Ocellar setae arising beside, or slightly in front of, anterior ocellus. Eye bare. Postpronotum restricted to the three main postpronotal setae. Dark stripes on either side of dorsocentral row of setae separated from one another by yellow tomentosity. Median and lateral stripes on either side of scutum separate from each other posteriorly. Postsutural dorsocentral setae 4. Anterior quadrant of anepisternum covered with long setae, none distinctly larger than the others or forming a row. Katepisternum with three setae, the middle one always the smallest. Vein R1 bare dorsally. Legs ranging from reddish brown to yellow tinged but overall dark. Coxae dark usually concolourous with remainder of leg. Ground colour of dorsal surface of abdomen dark to black. Ground colour of ventral surface of abdomen entirely black. Sex patches present on tergites 4 and 5. Ground colour of sex patches shiny black. Terminalia: surstylus bayonet shaped, posterodorsal half bare, apex bearing many stout apical spines, tip with strong inwardly 78 · Zootaxa 3858 (1) © 2014 Magnolia Press FLEMING ET AL. apical curve when viewed dorsally. Cerci rounded, apex with blunt, hooked tip, ventral surface haired, separation between cerci straight, up to 85% as long as surstylus. Lobe of sternite 5 small and pointed apically, inner margin covered in dense tomentosity appearing darker than surrounding cuticle, internal edge inwardly curved, single long apical seta emanating from apex. Hosts. Houghia pilosifrons has been reared 7 times, from a sample of 3500+ rain forest and dry forest Megalopygidae: Megalopyge lanata (Stoll) (1X), Podalia orsilocha Cramer (4X), and Trosia nigropunctigera Fletcher (2X). Holotype. ♂, CNC. Type locality: Costa Rica, Area de Conservación Guanacaste, Prov. Alajuela, Sector Rincon Rain Forest, Jacobo (10.94076°, -85.31170°), 461 m, 08/13/2009, Noe Castillo, DHJPAR0037559. Paratypes. 78 ♂, 80 ♀ (CNC) Costa Rica, Prov. Guanacaste, ACG database codes: DHJPAR0008822, DHJPAR0008823, DHJPAR0008824, DHJPAR0008825, DHJPAR0016102, DHJPAR0029682, 04-SRNP-40371, 03-SRNP-12380.1, 06-SRNP-7049, 95-SRNP-8077, 09-SRNP-69792, 08-SRNP-72108. Etymology. From the Latin adjective “pilosus”, meaning covered with hair, and the noun “frons”, meaning forehead, in reference to the denser small hairs entirely covering the fronto-orbital plates, thus a noun in apposition. Distribution. Costa Rica, ACG, Prov. Alajuela & Guanacaste, rain forest and dry forest, 260–645 m elevation. FIGURE 40. General morphology of Houghia pilosifrons sp. nov. a. Dorsal habitus. b. Frontal view of head. c. Lateral habitus. d. Posterior view of male terminalia. e. Lateral view of male terminalia. f. Ventral view of sternite 5 + segments 7+8. HOUGHIA OF ACG, COSTA RICA Zootaxa 3858 (1) © 2014 Magnolia Press · 79 Houghia romeroae Fleming & Wood, sp. nov. Figs. 2 a, 41 a–c Diagnosis. Houghia romeroae belongs to the H. blancoi group of species (H. blancoi, H. romeroae, H. chavarriae, H. marini, and H. matarritai). It has few distinctive features except in the male terminalia and the the presence of gold tomentosity only directly adjacent to the ocellar triangle, covering less than 25% of the fronto-orbital plate. The eye is bare and the antenna is entirely black. The ground colour of the abdomen is black, and sex patches are confined to tergites 4 and 5. The apex of the first flagellomere is shorter than the face by half the length of the pedicel. Gold tomentosity on the frons is diffuse and not well defined in area, but extends from vertex to the upper fronto-orbital setae (Fig. 41b), thus is more extensive than in H. blancoi. FIGURE 41. General morphology of Houghia romeroae sp. nov. a. Dorsal habitus. b. Frontal view of head. c. Lateral habitus. Description. Male. Antenna black. When viewed in profile, antenna arises distinctly above middle of eye. Length of first flagellomere almost extending to facial margin (usually shorter by less than length of pedicel). Facial ridge bare except for a few (usually 3–5) decumbent small setae above vibrissa. Palpus pale, usually distinctly yellowish. Postgena behind postoccipital row, above level of lower facial margin, with a small patch of few black setae. Parafacial silver. Colour of fronto-orbital plate gold only at vertex, adjacent to ocellar triangle, the remainder silver (up to 25% coverage). Surface of fronto-orbital plate almost bare. Ocellar triangle, when viewed from above appearing to be notched anteriorly. Diameter of anterior ocellus equal to, or greater than, diameter of base of adjacent ocellar seta. Ocellar setae arising beside, or slightly in front of, anterior ocellus. Eye bare. Postpronotum with 4 or 5 postpronotal setae. Dark stripes on either side of dorsocentral row of setae separated from one another by yellow tomentosity. Median and lateral stripes on either side of scutum separate from each other posteriorly. Postsutural dorsocentral setae 4. Anterior quadrant of anepisternum covered with short setae except for usually 3 to 5 distinctly larger setae. Katepisternum with three setae, the middle one always the smallest. Vein R1 bare dorsally. Legs ranging from reddish brown to yellow tinged but overall dark. Coxae dark usually concolourous with remainder of leg. Ground colour of dorsal surface of abdomen dark to black. Ground colour of ventral surface of abdomen entirely black. Sex patches present on tergites 4 and 5. Ground colour of sex patches shiny black. Terminalia (not illustrated due to poor quality of specimen): surstylus equilaterally oblong shaped, posterodorsal half bare, apex bearing apical spines along lower half only, tip with strong inwardly apical curve when viewed dorsally. Cerci rounded, apex with blunt, hooked tip, ventral surface haired, about as long as surstylus. Data for sternite 5 was unavailable, and due to the scarcity of this species could not be obtained at this time. Only one male remains undissected, and this specimen is designated as the holotype. 80 · Zootaxa 3858 (1) © 2014 Magnolia Press FLEMING ET AL. Hosts. Houghia romeroae has been reared only 6 times, from a sample of 74 wild-caught dry forest caterpillars of Lacosoma maldera Schaus (Mimallonidae). This fly was reared from no other species of caterpillar. Holotype. ♂, CNC. Type locality: Costa Rica, Area de Conservación Guanacaste, Prov. Guanacaste, Sector El Hacha, Estación Los Almendros (11.03226°, -85.52776°), 290 m, 11/25/2008, Lucia Ríos, DHJPAR0030201. Paratypes. 1 ♂, 3 ♀ (CNC) Costa Rica, Guanacaste, ACG database codes: DHJPAR0008132, DHJPAR0008123, DHJPAR0008142, DHJPAR0008117. Etymology. Houghia romeroae is dedicated to Luz Maria Romero of Liberia, Guanacaste, Costa Rica, in recognition of her extreme enthusiasm for helping the ACG parataxonomists that rear these tachinids to improve their caterpillar and parasitoid inventory abilities. Distribution. Costa Rica, ACG, Prov. Guanacaste, dry forest, 285–300 m elevation. Houghia sexmaculata Fleming & Wood, sp. nov. Figs. 1a, 4a, 8b, 42 a–e Diagnosis. The males of H. sexmaculata are characterized by the presence of dark gray to black palpus, 2 katepisternal setae, tergite 3 as well as tergites 4 and 5 with ventral sex patches, all on black integument, and dorsal surface of abdomen uniformly yellowish tomentose, with scarcely detectable dark apical bands. Females are distinctive in having a dark palpus and unmarked abdomens. Description. Male. Antenna black. When viewed in profile, antenna arises approximately at level of middle of eye. Length of first flagellomere almost extending to facial margin (usually shorter by less than length of pedicel). Facial ridge bare except for a few (usually 3–5) decumbent small setae above vibrissa. Palpus dark, gray to black. Postgena behind postoccipital row, above level of lower facial margin, with a small patch of few black setae. Parafacial silver. Colour of fronto-orbital plate gold only at vertex, adjacent to ocellar triangle, the remainder silver (up to 25% coverage). Surface of fronto-orbital plate almost bare. Ocellar triangle, when viewed from above appearing rounded anteriorly. Diameter of anterior ocellus less than diameter of base of adjacent ocellar seta. Ocellar setae arising behind anterior ocellus but closer to anterior than to posterior ocelli. Eye bare. Postpronotum with 4 or 5 postpronotal setae. Dark stripes on either side of dorsocentral row of setae separated from one another by yellow tomentosity. Median and lateral stripes on either side of scutum separate from each other posteriorly. Postsutural dorsocentral setae 3. Anterior quadrant of anepisternum covered with short setae except for usually 3 to 5 distinctly larger setae. Katepisternum with two setae. Vein R1 bare dorsally. Legs ranging from reddish brown to yellow tinged but overall dark. Coxae dark usually concolourous with remainder of leg. Ground colour of dorsal surface of abdomen dark to black. Ground colour of ventral surface of abdomen entirely black. Sex patches present on tergite 3, as well as on tergites 4 and 5 (though sometimes small or absent on 5). Ground colour of sex patches shiny black. Terminalia: surstylus equilaterally oblong, posterodorsal half bare, apex bearing many stout apical spines, tip with a slight outward curve. Cerci rounded, apex with straight tip, ventral surface bare, separation between cerci straight, up to 85% as long as surstylus. There is insufficient data to comment on sternite 5, as it appears it may have been damaged on dissection. However, this data not necessary for differential diagnosis of this species. Hosts. To date, H. sexmaculata has been reared from 6 of the 10 species of Chlosyne Butler (Nymphalidae) that occur in ACG dry forest and rain forest, with 1,474 wild-caught caterpillars yielding 63 rearings of this species; it has been reared only from Chlosyne. Holotype. ♂, CNC. Type locality: Costa Rica, Area de Conservación Guanacaste, Prov. Guanacaste, Sector El Hacha, Finca Araya (11.01541°, -85.51125°), 295 m, 10/09/2002, Manuel Pereira, DHJPAR0019227. Paratypes. 20 ♂, 25 ♀ (CNC) Costa Rica, Prov. Guanacaste, ACG database codes: DHJPAR0016637, DHJPAR0030155, DHJPAR0029863, DHJPAR0019228, DHJPAR0019230, DHJPAR0019231, DHJPAR0030152, DHJPAR0019237, DHJPAR0019236, DHJPAR0030161, DHJPAR0011489, DHJPAR0030177, DHJPAR0019239, DHJPAR0019241, DHJPAR0029868, DHJPAR0029819, DHJPAR0029874, DHJPAR0030174, DHJPAR0029869, DHJPAR0029871, DHJPAR0029872, DHJPAR0019235, DHJPAR0030176, DHJPAR0029867, DHJPAR0019229, DHJPAR0030171, DHJPAR0019248, DHJPAR0019234, DHJPAR0029723, DHJPAR0019244, DHJPAR0019233, DHJPAR0019240, DHJPAR0019245, DHJPAR0030179, DHJPAR0030157, DHJPAR0019225, DHJPAR0029873, DHJPAR0030151, DHJPAR0029861, DHJPAR0030170, DHJPAR0019242, DHJPAR0029227, DHJPAR0011486, HOUGHIA OF ACG, COSTA RICA Zootaxa 3858 (1) © 2014 Magnolia Press · 81 DHJPAR0019232, DHJPAR0016335; and not databased or barcoded, 1 ♀ COSTA RICA, Gste, 5 km.N.Cañas, 9.XII.1990, M.Polak & M.Wood. Etymology. From the Latin “sex” meaning six, and the adjective “maculatus”, meaning spotted or marked, thus meaning six-spotted in reference to the three pairs of sex patches on black integument on the undersurface of the abdomen; other species (H. gracilis and H. graciloides) have sex patches on tergite 3, but on a reddish integument, and are less prominent. Distribution. Costa Rica, ACG, Prov. Guanacaste, dry forest and rain forest, 20–540 m elevation. FIGURE 42. General morphology of Houghia sexmaculata sp. nov. a. Dorsal habitus. b. Frontal view of head. c. Lateral habitus. d. Posterior view of male terminalia. e. Lateral view of male terminalia. Houghia spathulata Fleming & Wood, sp. nov. Fig. 43 a–f Diagnosis. Although sequence data suggests that it is more closely related to H. aerata, H. spathulata closely resembles members of the cluster of species surrounding H. blancoi (H. blancoi, H. romeroae, H. chavarriae, H. marini, and H. matarritai). It is distinguished from the other species within that cluster by, by its barcode, and by the elongate, straight-sided surstylus (Fig. 43e) that extends far beyond the apex of the cerci. Description. Male. Antenna black. When viewed in profile, antenna arises approximately at level of middle of eye. Length of first flagellomere almost extending to facial margin (usually shorter by less than length of pedicel). 82 · Zootaxa 3858 (1) © 2014 Magnolia Press FLEMING ET AL. Facial ridge bare except for a few (usually 3–5) decumbent small setae above vibrissa. Palpus pale, usually distinctly yellowish. Postgena behind postoccipital row, above level of lower facial margin, with a small patch of few black setae or extensively covered with setae, and fine hairs. Parafacial silver. Colour of fronto-orbital plate pale brassy to gold on its entire length from vertex to base of antenna (more than 50% coverage). Surface of frontoorbital plate covered with small recumbent hairs, especially near margin of eye. Ocellar triangle, when viewed from above, appearing rounded anteriorly. Diameter of anterior ocellus equal to, or greater than, diameter of base of adjacent ocellar seta. Ocellar setae arising beside, or slightly in front of, anterior ocellus. Eye with inconspicuous hairs. Postpronotum with 4 or 5 postpronotal setae. Dark stripes on either side of dorsocentral row of setae separated from one another by yellow tomentosity. Median and lateral stripes on either side of scutum separate from each other posteriorly. Postsutural dorsocentral setae 4. Anterior quadrant of anepisternum covered with long setae, none distinctly larger than the others or forming a row. Katepisternum bearing three setae, the middle one always the smallest. Vein R1 bare dorsally. Legs ranging from reddish brown to yellow tinged but overall dark. Coxae dark, usually concolourous with remainder of leg. Ground colour of dorsal surface of abdomen dark to black. Ground colour of ventral surface of abdomen entirely black. Sex patches present on tergites 4 and 5. Ground colour of sex patches shiny black. Terminalia: surstylus equilaterally oblong shaped, posterodorsal half bare, apex bearing many stout apical spines, tip with light inwardly apical curve when viewed dorsally. Cerci sharply pointed or distinctly tapered, apex with blunt, hooked tip, ventral surface haired, separation between cerci deep narrow V shape, up to 85% as long as surstylus. Lobe of sternite 5 small and pointed apically, inner margin covered in dense tomentosity appearing darker than surrounding cuticle, internal edge inwardly curved, apical seta absent. FIGURE 43. General morphology of Houghia spathulata sp. nov. a. Dorsal habitus. b. Frontal view of head. c. Lateral habitus. d. Posterior view of male terminalia. e. Lateral view of male terminalia. f. Ventral view of sternite 5 + segments 7+8. HOUGHIA OF ACG, COSTA RICA Zootaxa 3858 (1) © 2014 Magnolia Press · 83 Hosts. Houghia spathulata has been reared only 3 times, among 50000+ caterpillars of Notodontidae, Noctuidae and Geometridae; owing to the variety of hosts and its DNA barcode variability, this morphology-based species name is likely to be found to cover two species of tachinid when a larger sample is available. Holotype. ♂, CNC. Type locality: Costa Rica, Area de Conservación Guanacaste, Prov. Guanacaste, Sector Santa Elena, Vado Descanso (10.86336°, -85.72535°), 70 m, 01/05/2003, Guillermo Pereira, DHJPAR0008830. Paratypes. 4 ♂ (CNC) Costa Rica, Prov. Guanacaste, ACG database codes: DHJPAR0048673, DHJPAR0040647, 03-SRNP-25. Etymology. An adjective derived from the Latin noun “spatha”, a broad, two-edged sword without a point, meaning “spatha-shaped” or like a spatha, referring to the shape of the surstylus. Distribution. Costa Rica, ACG, Prov. Alajuela & Guanacaste, rain forest and dry forest, 70–380 m elevation. Houghia triangularis Fleming and Wood, sp. nov. Fig. 44 a–f Diagnosis. One of a group of seven closely related species in Guanacaste, the H. crypta group in which the second postsutural dorsocentral seta is missing, leaving a gap between first and third setae (H. omissa, H. confinis, H. triangularis, H. destituta, H. crypta, H. biseriata, H. bivittata). Compared to the other species, the anterior portion of the ocellar triangle of this species is more sharply pointed apically, more distinctly gold pollinose, and is visible in both side and frontal views (Fig. 44b). Description. Male. Antenna black. When viewed in profile, antenna arises approximately at level of middle of eye. Length of first flagellomere almost extending to facial margin (usually shorter by less than length of pedicel). Facial ridge bare except for a few (usually 3–5) decumbent small setae above vibrissa. Palpus pale, usually distinctly yellowish. Postgena behind postoccipital row, above level of lower facial margin, with a small patch of few black setae. Parafacial silver. Colour of fronto-orbital plate gold only at vertex, adjacent to ocellar triangle, the remainder silver (up to 25% coverage). Surface of fronto-orbital plate almost bare. Ocellar triangle when viewed from above appearing notched anteriorly. Diameter of anterior ocellus equal to, or greater than, diameter of base of adjacent ocellar seta. Ocellar setae arising beside, or slightly in front of, anterior ocellus. Eye bare. Postpronotum restricted to the three main postpronotal setae. Dark stripes on either side of dorsocentral row of setae, separated from one another by yellow tomentosity. Median and lateral stripes on either side of scutum separate from each other posteriorly. Postsutural dorsocentral setae 3, with a gap between first and third seta. Anterior quadrant of anepisternum covered with short setae except for usually 3 to 5 distinctly larger setae. Katepisternum bearing three setae, the middle one always the smallest. Vein R1 bare dorsally. Legs ranging from reddish brown to yellow tinged but overall dark. Coxae dark, usually concolourous with remainder of leg. Ground colour of dorsal surface of abdomen dark to black. Ground colour of ventral surface of abdomen entirely black. Sex patches present on tergites 4 and 5. Ground colour of sex patches shiny black. Terminalia: surstylus equilaterally oblong shaped, posterodorsal half haired, apex bearing many stout apical spines, tip with strong inwardly apical curve when viewed dorsally. Cerci sharply pointed or distinctly tapered, apex with blunt, hooked tip, ventral surface haired, separation between cerci deep narrow V shape, up to 85% as long as surstylus. Lobe of sternite 5 large and rounded apically, inner margin covered in dense tomentosity appearing darker than surrounding cuticle, internal edge inwardly curved, apical seta absent. Hosts. Houghia triangularis has been reared only 6 times, from 3 species of satyrine Nymphalidae feeding on grasses (Poaceae), among a sample of 800+ caterpillars of these species. Holotype. ♂, CNC. Type locality: Costa Rica, Area de Conservación Guanacaste, Prov. Guanacaste, Sector Pitilla, Ingas (11.00311°, -85.42041°), 580 m, 12/05/2011, Freddy Quesada, DHJPAR0048572. Paratypes. 3 ♂, 2 ♀ (CNC) Costa Rica, Prov. Guanacaste, ACG database codes: DHJPAR0042571, DHJPAR0027975, DHJPAR0048458, DHJPAR0015258, DHJPAR0046961. Etymology. From the Latin adjective “triangulus”, meaning three-cornered, or triangular, referring to the pointed rather than notched anterior angle of the ocellar triangle. Distribution. Costa Rica, ACG, Prov. Guanacaste, rain forest and dry forest, 290–675 m elevation. 84 · Zootaxa 3858 (1) © 2014 Magnolia Press FLEMING ET AL. FIGURE 44. General morphology of Houghia triangularis sp. nov. a. Dorsal habitus. b. Frontal view of head. c. Lateral habitus. d. Posterior view of male terminalia. e. Lateral view of male terminalia. f. Ventral view of sternite 5 + segments 7+8. Houghia velutina Fleming & Wood sp. nov. Fig. 45 a–f Diagnosis. Similar to H. latigena, and part of the H. latigena group of related species in which the gena is 1/10 head height (H. longicercus, H. latigena, H. latilobus, and H. velutina). The height of the V-shaped apical notch of the cerci, in posterior view, is only 1/3 of the height of the cerci, as in H. latigena, but differs from that species by being more evenly tapered to its apex rather than being slightly constricted at midlength (compare Figs. 27d, 28d, 30d). Description. Male. Antenna black. When viewed in profile, antenna arises approximately at level of middle of eye. Length of first flagellomere extending to facial margin. Facial ridge bare except for a few (usually 3–5) decumbent small setae above vibrissa. Palpus pale, usually distinctly yellowish. Postgena behind postoccipital row, above level of lower facial margin, with a small patch of few black setae. Parafacial silver. Colour of fronto-orbital plate pale brassy to gold on its entire length from vertex to base of antenna (more than 50% coverage). Surface of fronto-orbital plate covered with small recumbent hairs, especially near margin of eye. Ocellar triangle, when viewed from above appearing rounded anteriorly. Diameter of anterior ocellus less than diameter of base of adjacent ocellar seta. Ocellar setae arising behind anterior ocellus but closer to anterior than to posterior ocelli. Eye HOUGHIA OF ACG, COSTA RICA Zootaxa 3858 (1) © 2014 Magnolia Press · 85 bare. Postpronotum restricted to the three main postpronotal setae. Dark stripes on either side of dorsocentral row of setae separated from one another by yellow tomentosity. Median and lateral stripes on either side of scutum united to one another posteriorly. Postsutural dorsocentral setae 4. Anterior quadrant of anepisternum covered with short setae except for usually 3 to 5 distinctly larger setae. Katepisternum with three setae, the middle one always the smallest. Vein R1 bare dorsally. Legs ranging from reddish brown to yellow tinged but overall dark. Coxae dark, usually concolourous with remainder of leg. Ground colour of dorsal surface of abdomen dark to black. Ground colour of ventral surface of abdomen entirely black. Sex patches present on tergites 4 and 5. Ground colour of sex patches shiny black. Terminalia: surstylus wedge shaped, posterodorsal half haired, apex devoid of apical spines, tip straight. Cerci rounded, apex with blunted hook tip, ventral surface haired, separation between forming a deep narrow V shape, about as long as surstylus. Lobe of sternite 5 large and rounded apically, inner margin covered in dense tomentosity appearing darker than surrounding cuticle, internal edge straight, single long apical seta emanating from apex. Hosts. This rare fly has been reared only two times, and from 1081 caterpillars of Yanguna cosyra Druce (Pyrginae, Hesperiidae) feeding on Clusia L. and Chrysochlamys Poeppig (Clusiaceae), in ACG rain forest. Holotype. ♂, CNC. Type locality: Costa Rica, Area de Conservación Guanacaste, Prov. Alajuela, Sector Rincon Rain Forest, Jacobo (10.94076°, -85.31770°), 461 m, 06/17/2009, Keiner Aragon, DHJPAR0035853. Paratypes. 5 ♂, 2 ♀ (CNC) Costa Rica, Prov. Alajuela, ACG database codes: DHJPAR0020985, 09-SRNP69334, 07-SRNP-41564. FIGURE 45. General morphology of Houghia velutina sp. nov. a. Dorsal habitus. b. Frontal view of head. c. Lateral habitus. d. Posterior view of male terminalia. e. Lateral view of male terminalia. f. Ventral view of sternite 5 + segments 7+8. 86 · Zootaxa 3858 (1) © 2014 Magnolia Press FLEMING ET AL. Etymology. An adjective derived from the Latin noun “vellus”, meaning fleece, or wool when shorn off, referring to the velvety appearance of the sex patches. Distribution. Costa Rica, ACG, Prov. Alajuela, rain forest, 405–461 m elevation. Acknowledgments We gratefully acknowledge the unflagging support of the team of ACG parataxonomists (Janzen et al 2009, Janzen & Hallwachs 2011) who found and reared the specimens used in this study, and the team of biodiversity managers who protect and manage the ACG forests that host these tachinids and their caterpillar hosts. The study has been supported by U.S. National Science Foundation grants BSR 9024770 and DEB 9306296, 9400829, 9705072, 0072730, 0515699, and grants from the Wege Foundation, International Conservation Fund of Canada, Jessie B. Cox Charitable Trust, Blue Moon Fund, Guanacaste Dry Forest Conservation Fund, Area de Conservación Guanacaste, Permian Global, and University of Pennsylvania (DHJ & WH). This study has been supported by the Government of Canada through its ongoing support of the Canadian National Collection, Genome Canada, the Biodiversity Institute of Ontario, and the Ontario Genomics Institute (2008-0GI-ICI-03) (MAS), and by a Discovery Grant from Natural Sciences and Engineering Research Council of Canada (MAS). The authors also wish to acknowledge the efforts and time of the editor and reviewers, who provided insights and suggestions allowing us to make our work better. Literature cited Aldrich, J.M. (1927) Redescription of types of American muscoid flies in the collection of the Vienna Natural History Museum with incidental notes. Proceedings of the United States National Museum, 72 (Art. 7) [= No. 2703] [1928], 1–35. http://dx.doi.org/10.5479/si.00963801.72-2703.1 Arnaud, P.H., Jr. (1963) Types of the Tachinidae (Diptera) in the American Museum of Natural History. Bulletin of the American Museum of Natural History, 125, 101–137. 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