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1 Zootaxa 3774 (1): Copyright 2014 Magnolia Press Article The Neotropical species of Xanthopimpla Saussure (Hymenoptera: Ichneumonidae: Pimplinae) ISSN (print edition) ZOOTAXA ISSN (online edition) ISRRAEL C. GÓMEZ 1,5, ILARI E. SÄÄKSJÄRVI 1, GAVIN R. BROAD 2, LIISA PUHAKKA 1, CAROL CASTILLO 1, CARLOS PEÑA 3 & DIEGO G. PÁDUA 4 1 Zoological Museum, Section of Biodiversity and Environmental Sciences, Department of Biology, FI University of Turku, Finland 2 Department of Life Sciences, The Natural History Museum, London,UK 3 Laboratory of Genetics, Department of Biology, FI University of Turku, Finland 4 Programa de Pós Graduação em Entomologia, Instituto Nacional de Pesquisas da Amazônia INPA, Brazil 5 Corresponding author. isrrael.gomez@yahoo.com Abstract Xanthopimpla Saussure, 1892 is one of the largest and best studied genera of the family Ichneumonidae. It is most species rich in the Oriental and Afrotropical regions with only a few species occurring in Central and South America. The present study reviews the Neotropical species of the genus including descriptions of four new species from Amazonia and Northeast South America. We define a new species group: the amazonica species-group, to accommodate the following five species: X. amazonica Gómez, Sääksjärvi & Veijalainen, X. guianensis Gómez & Sääksjärvi sp. n., X. jussilai Veijalainen, Sääksjärvi & Broad, X. pucallpensis Gómez & Sääksjärvi sp. n. and X. vidali Gómez sp. n. The aurita species-group, which had hitherto been regarded as the only species-group in the Neotropical region, is currently represented by five species: X. allpahuaya Gómez & Sääksjärvi sp. n., X. aurita Krieger, X. craspedoptera Krieger, X. rhabdomera Townes and X. spiloptera Krieger. The Andean species X. peruana Krieger is established as an unplaced species outside of the amazonica and aurita species-groups. A key to Neotropical species-groups and species of Xanthopimpla is provided. Xanthopimpla aurita is recorded for the first time from Ecuador and Colombia and its extensive distribution is discussed. Xanthopimpla amazonica, X. craspedoptera and X. jussilai are recorded for the first time from Brazil; X. amazonica is recorded for the first time from French Guiana; X. spiloptera is recorded for the first time from French Guiana and Peru, and X. rhabdomera is recorded for the first time from Peru. Key words: taxonomy, parasitoids, idiobiont, new species, Amazonia, Peru, Ecuador, French Guiana, Brazil Resumen Xanthopimpla Saussure, 1892 es uno de los géneros más grandes y mejor estudiados de la familia Ichneumonidae. Este género es más rico en especies en las regiones Oriental y Afrotropical con solo algunas especies presentes en Centroamérica y Sudamérica. En este estudio, examinamos las especies Neotropicales del género incluyendo descripciones de cuatro nuevas especies encontradas en la Amazonia y la parte Noreste de Sudamerica. Se define un grupo nuevo de especies, el grupo amazonica, para acomodar las siguientes cinco especies: X. amazonica Gómez, Sääksjärvi & Veijalainen, X. guianensis Gómez & Sääksjärvi sp. n., X. jussilai Veijalainen, Sääksjärvi & Broad, X. pucallpensis Gómez & Sääksjärvi sp. n. y X. vidali Gómez sp. n. El grupo de especies aurita, que fue anteriormente considerado como el único grupo de especies para la región Neotropical, está actualmente representado por cinco especies X. allpahuaya Gómez & Sääksjärvi sp. n., X. aurita Krieger, X. craspedoptera Krieger, X. rhabdomera Townes y X. spiloptera Krieger. La especie andina X. peruana Krieger es establecida independientemente fuera de los grupos amazonica y aurita. Se presenta una clave actualizada para los grupos de especies y especies Neotropicales de Xanthopimpla. La especie Xanthopimpla aurita es registrada por primera vez de Ecuador y Colombia y se discute su amplia distribución. Xanthopimpla amazonica, X. craspedoptera y X. jussilai son registradas por primera vez de Brazil; X. amazonica es registrada por primera vez en Guayana Francesa; X. spiloptera es registrada por primera vez en Guayana Francesa y Perú y X. rhabdomera es registrada por primera vez de Perú. Palabras clave: taxonomía, parasitoide, idiobionte, nuevas especies, Amazonia, Perú, Ecuador, Guayana Francesa, Brasil. Accepted by J. T Jennings: 21 Jan. 2014; published: 4 Mar

2 Introduction The parasitoid wasp family Ichneumonidae is one of the most species rich components of insect life in most terrestrial ecosystems (Gauld 1991; Sääksjärvi et al. 2004). However, only a few detailed ichneumonid inventories have been conducted in rainforests of the Neotropical region. In this region, most studies have concentrated on Central America, in particular Costa Rica, where the late Ian D. Gauld with his collaborators monographed many of the subfamilies (Gauld 1988; 1991; 2000; Gauld et al. 1997; 1998; 2002). These revisionary works hugely improved our knowledge of the Neotropical fauna. Although the South American ichneumonid fauna is generally far more poorly known than the Central American fauna, over the last 15 years, some highly diverse regions of western South America have been sampled in more detail. Sampling has mainly concentrated on the tropical Andes (e.g. Palacio et al. 2007; Castillo et al. 2011) and Western Amazonian areas (e.g. Sääksjärvi et al. 2003; 2004; Gómez et al. 2009; Palacio et al. 2010; Broad et al. 2011). These studies have revealed a plethora of new species and genera from the Andean and Amazonian interface. Based on this evidence, and considering that most of the Neotropical region is still poorly studied for parasitoids, the ichneumonid fauna in this region is certainly one of the richest on Earth. Following the taxonomic work of, especially, Gauld (Gauld 1991; Gauld et al., 1998; 2002) and Charles Porter (Porter 1970 a,b) the ichneumonid subfamily Pimplinae is now one of the best-known groups of the family in the Neotropical region. Species of Xanthopimpla Saussure are among the most conspicuous and abundant tropical components of pimplines, predominantly bright yellow, moderately large and rather easily defined and recognized by several reliable morphological characters. For a long time this genus has caught the attention of various entomologists (e.g. Thunberg 1822; Saussure 1892; Tosquinet 1896; Krieger 1899; Krieger 1914; Townes 1969; Townes & Chiu 1970; Gómez et al. 2009, Pham et al. 2011) becoming to date the most species-rich pimpline genus and one of the best studied genera of the family Ichneumonidae. Xanthopimpla species are idiobiont endoparasitoids of Lepidoptera pupae (Gauld 1991). For this reason, several species have been utilized in biological control of economically important agricultural pests (Hailemichael et al. 1994; Gitau et al. 2007; Dum et al. 2011). Many species are abundant in tropical areas, especially in tropical Asia where most of the species occur (Townes & Chiu 1970; Pham et al. 2011). Surprisingly, only a few Xanthopimpla species have been described from the Neotropical region (Townes 1969; Gómez et al. 2009). It is, however, important to consider that many tropical pimplines are hard to collect even during long-term field inventories (Sääksjärvi et al. 2004) and the rare species usually remain undiscovered by short-term sampling. The aim of the present work is to revise the Neotropical Xanthopimpla. We describe four new species and define a new species-group to accommodate several South American species which cannot be accommodated within any of the numerous existing species-groups (most of which are strictly Old World). In addition, we illustrate and provide identification keys to the Neotropical species-groups and species. Material and methods This study is based mainly on the ichneumonid samples collected during our long-term western Amazonian ichneumonid survey in the years 1998, 2000 (Sääksjärvi et al. 2004), 2008 (Gómez et al. 2009) and 2011 (unpublished data). During this survey we have utilized different sampling methods (mainly Malaise trapping but also sweep netting, rearing, light trapping and yellow pan trapping) in Allpahuayo-Mishana National Reserve (Department of Loreto, Peru) and in Los Amigos Conservation concession (Department of Madre de Dios, Peru). However, Malaise trapping has been the only successful method in sampling Xanthopimpla. In addition, we have studied a large number of samples collected by canopy fogging in Ecuadorean Amazonia (by Terry Erwin and his team) and Malaise samples collected in tropical Andean forests (Castillo et al. 2011). Over the last few years we have also studied a large number of Malaise samples collected from the tropical forests of French Guiana (by the Société Entomologique Antilles Guyane (SEAG) team; ichneumonid samples coordinated by Yves Braet) and Central America (LLAMA project, led by John Longino). We have also studied a number of specimens deposited in the Instituto Nacional de Pesquisas da Amazônia, Brazil (INPA) and the Museu de Zoologia da Universidade de São Paulo, Brazil (MZUSP). 58 Zootaxa 3774 (1) 2014 Magnolia Press GÓMEZ ET AL.

3 In addition to the extensive new material listed above, we studied the Neotropical specimens of Xanthopimpla deposited in the Natural History Museum, London, UK (BMNH) and the Swedish Museum of Natural History, Stockholm, Sweden (NHRS). Paratypes of X. rhabdomera Townes deposited in the Canadian National Collection of Insects (CNC) were studied using photos provided by Andrew Bennet. Morphological terminology and style of descriptions follow Gauld (1991) and Gómez et al. (2009). The identification key for the aurita species-group is modified from that of Gómez et al. (2009). Specimens are deposited in the following collections: the Zoological Museum, Section of Biodiversity and Environmental Science, Department of Biology, University of Turku, Finland (ZMUT); the Smithsonian Institution, Washington, DC, USA (NMNH); the Natural History Museum, University of San Marcos, Lima, Peru (UNSM); and the Instituto Nacional de Pesquisas da Amazônia, Brazil (INPA). All observations were made using Olympus SZX10 and SZ40 stereomicroscopes. Digital photos were taken using an Olympus SZX16 stereomicroscope attached to an Olympus E520 digital camera and combined using the software Deep Focus 3.1 and Quick PHOTO CAMERA 2.3. Taxonomy The following keys work best for females. Males of many species are unknown. For this reason, the identification of male specimens should always be confirmed by comparing them to female specimens and descriptions. A few specimens of many Neotropical species have dark-marked metasoma 5 +; identification of these specimens is more difficult and all characters should be checked. Key to the species-groups and unplaced species of Neotropical Xanthopimpla 1 Occipital carina complete and conspicuously developed as a thin, high flange (Fig. 1b), propodeal spiracle joined to pleural carina by a short carina (Fig. 1d) X. amazonica species-group - Occipital carina incomplete to complete, never developed into a flange (Fig. 1a), propodeal spiracle not joining pleural carina by a short carina (Fig. 1d) Propodeum with only posterior transverse carina present, triangular crests at mesoscutum small (Fig. 1c), body almost entirely bright lemon yellow (Fig. 5d) X. peruana Krieger - Propodeum with posterior transverse and, at least partially, anterior transverse carina present (Fig. 6a), triangular crests at anterior of mesoscutum highly elevated, body not uniformly bright lemon yellow X. aurita species-group Key to species of the amazonica species-group 1. Mesosternum yellow, ovipositor sheath more than 1.2 times as long as hind tibia, ovipositor tip strongly decurved (Fig. 2e) X. vidali Gómez sp. n. - Mesosternum blackish or at least with blackish spots, ovipositor sheath less than or equal to 1.2 times as long as hind tibia, ovipositor tip slightly decurved (Fig. 2a) Lateral longitudinal carina strongly defined and complete throughout propodeum (4b), posterior tergites of metasoma (4+) blackish X. guianensis Gómez & Sääksjärvi sp. n. - Lateral longitudinal carina of propodeum very weakly defined or incomplete medially, posterior tergites of metasoma (4+) usually yellowish Petiolar area divided by a weak longitudinal carina that starts in the center of posterior transverse carina (Fig. 3a), posterior transverse carina bowed centrally X. amazonica Gómez, Sääksjärvi & Veijalainen - Petiolar area not divided by a weak longitudinal carina that starts in the center of posterior transverse carina, posterior transverse carina not bowed centrally (Fig. 3e) Propodeum with area dentipara defined (Fig. 3c), tergite 1 about 1.2 times as long as posteriorly broad X. jussilai Veijalainen, Sääksjärvi & Broad - Propodeum with area dentipara not defined (Fig. 3d), tergite 1 about 1.3 times as long as posteriorly broad X. pucallpensis Gómez & Sääksjärvi sp. n. Key to species of the aurita species-group 1. Posterior transverse carina bowed centrally (Fig. 6b), ovipositor sheath about times as long as hind tibia X. aurita Krieger - Posterior transverse carina not as above, ovipositor sheath less than 0.8 times as long as hind tibia THE NEOTROPICAL SPECIES OF XANTHOPIMPLA Zootaxa 3774 (1) 2014 Magnolia Press 59

4 2. Submetapleural carina with a small flange anteriorly (Fig. 7c), with black spots close to propodeal spiracle and ventrally on hind coxa x. craspedoptera Krieger - Submetapleural carina without a small flange anteriorly, without black spots surrounding the propodeal spiracle Propodeum with lateromedian longitudinal carina weak, only present anteriorly (Figs. 6e, 7e), ovipositor sheath entirely blackish, fifth hind tarsal segment blackish x. rhabdomera Townes - Propodeum with lateromedian longitudinal carina clearly defined, ovipositor sheath not entirely blackish, fifth hind tarsal segment yellowish Occipital carina complete, metasomal tergites with black bands, posterior transverse carina of propodeum angulate (Fig. 6f) X. spiloptera Krieger - Occipital carina incomplete dorsally, metasomal tergites without black bands, posterior transverse carina of propodeum rounded (Fig. 6a) X. allpahuaya Gómez & Sääksjärvi sp. n. The amazonica species-group (Figs. 1b, 1e, 1h) Diagnosis. The amazonica species-group can be distinguished from all other New and Old world species-groups of Xanthopimpla by the combination of the following characters: 1) occipital carina developed into a thin, high flange (Fig. 1b); 2) propodeal spiracle joined to the pleural carina by a short carina (Fig. 1e); and 3) first tergite with lateromedian carinae anteriorly highly elevated (Fig. 1h). At least the first two characters seem to be unique to the amazonica species-group. Remarks. The species of the group are restricted to South America and have been found mainly in the vast Amazonian lowland rainforest area (Fig. 8). All species are rare and we have only seen a few specimens. Xanthopimpla amazonica Gómez, Sääksjärvi & Veijalainen 2009 (Figs. 2a, 3a, 4a) Original description. See Gómez et al. (2009). Diagnosis. This species can be distinguished from all other Neotropical Xanthopimpla by the combination of the following characters: 1) occipital carina complete and expanded into a high thin flange; 2) propodeum with petiolar area divided by a longitudinal carina that starts in the center of the posterior transverse carina; 3) posterior transverse carina bowed in the center; and 4) ovipositor sheath about times the length of the hind tibia. Distribution. Brazil, French Guiana, Peru. New material examined. Brazil: Male. (INPA) Dept. of Maranhão, Açailândia S, W, XII Rafael, Oliveira & Vidal leg. Malaise trap. French Guiana: Female (ZMUT) Patawa. Seag leg. II Notes. The species is here recorded for the first time outside Peru (see Gómez et al. 2009). The Brazilian male is similar to the Peruvian female specimen. The French Guianan female is similar to the Peruvian female but has tergites 5+ dark brownish and a longer ovipositor (about 1.1 times as long as hind tibia whereas in the Peruvian female the same index is about 0.8). Xanthopimpla amazonica seems to be rare but widely distributed in tropical northern South America. Xanthopimpla guianensis Gómez & Sääksjärvi sp. n. (Figs. 2b, 3b, 4b) Type material. Holotype female (ZMUT). French Guiana, Kourou, Seag leg. X Paratypes: Male. (ZMUT) French Guiana, Kourou, Seag leg. X Male (BMNH). French Guiana, Kourou, Seag leg. III Zootaxa 3774 (1) 2014 Magnolia Press GÓMEZ ET AL.

5 FIGURE 1. Comparison between the aurita species-group (left), the amazonica species-group (center), and the unplaced species X. peruana (right). (a) Xanthopimpla aurita; (b) X. vidali; (c) X. peruana; (d) X. aurita; (e) X. guianensis; (f) X. peruana; (g) X. allpahuaya; (h) X. guianensis; and (i) X. peruana. FIGURE 2. Habitus of the amazonica species-group (female holotypes). (a) Xanthopimpla amazonica; (b) X. guianensis; (c) X. jussilai; (d) X. pucallpensis; (e) X. vidali. THE NEOTROPICAL SPECIES OF XANTHOPIMPLA Zootaxa 3774 (1) 2014 Magnolia Press 61

6 FIGURE 3. Propodeum, dorsal view, of the amazonica species-group (female holotypes). (a) Xanthopimpla amazonica; (b) X. guianensis; (c) X. jussilai; (d) X. pucallpensis; (e) X. vidali. Female (Fig. 2b). Head in dorsal view moderately short, with genae evenly narrowed behind eyes; frons biconcave; posterior ocellus separated from eye by about 0.5 times its diameter; occipital carina complete, ventrally, laterally and dorsally expanded into a very high thin flange (Fig. 1b); clypeus relatively flat, basally not clearly separated from face; clypeal apex truncate; malar space about 0.15 times as long as basal mandibular width; face polished and pubescent, about 1.0 times as wide as medially high. Pronotum with apical edge strongly reflexed and raised, overlapping propleura. Mesoscutum pubescent, with notauli strongly marked anteriorly, bounded anteriorly by high triangular crests; scutellum pubescent and convex, laterally with high carinae. Mesopleuron polished, with ventral part densely pubescent; epicnemial carina reaching well above level of lower corner of pronotum, ventrally raised. Metapleuron convex, smooth; submetapleural carina sharp but low, extending back to insertion of hind coxa. Propodeum in profile declivous (Fig. 4b); posterior transverse carina complete and slightly curved; lateromedian longitudinal carinae present defining area superomedia (Fig. 3b); lateral longitudinal carina present; pleural carina complete. Fore wing length about 12 mm; areolet complete; vein Rs slightly sinuous. Tergite 1 of metasoma about 1.2 times as long as posteriorly broad, with lateral longitudinal and lateromedian longitudinal carinae strongly developed anteriorly; tergite 2 about 0.7 times as long as posteriorly broad, with a more or less rhombic raised central area. Claws of hind leg large, without a basal lobe and with four strong setae basally. Ovipositor sheath 0.8 times as long as hind tibia; apex of ovipositor evenly tapered and slightly decurved. Coloration. Yellow with the following areas blackish: transverse bands across mesoscutum centrally and posteriorly, scuto-scutellar groove, anterior part of propodeum, interocellar area, stripes leading from posterior ocelli to dorso-lateral small patches on occipital area, small patch on mesosternum. Tergites brownish with posterior margins of tergites 1 3 yellow, posterior tergites 4+ completely blackish. Scape yellow in frontal view and brown dorsally. Ovipositor sheath proximally brown, infuscate distally. Wings slightly yellowish, pterostigma yellowish. Male. Similar to female. Diagnosis. This species can be distinguished from all other Neotropical Xanthopimpla by the combination of the following characters: 1) occipital carina complete and expanded into a high thin flange; 2) propodeum with complete and exceptionally strong carination; 3) tergite 1 about 1.2 times as long as wide; and 4) posterior ocellus rather small, separated from eye by about 0.5 times its diameter. Xanthopimpla guianensis resembles X. 62 Zootaxa 3774 (1) 2014 Magnolia Press GÓMEZ ET AL.

7 pucallpensis sp. n. However, its smaller ocellus, defined area dentipara, darker metasoma, and more robust body differentiate this species from X. pucallpensis sp. n. Biological notes. Nothing is known about the hosts of this species. Etymology. The name guianensis refers to French Guiana. This is the first Xanthopimpla species described from this country. Distribution. French Guiana. Xanthopimpla jussilai Veijalainen, Sääksjärvi & Broad 2009 (Figs. 2c, 3c, 4c) Original description. See Gómez et al. (2009). Diagnosis. This species can be distinguished from all other Neotropical Xanthopimpla by the combination of the following characters: 1) occipital carina complete and expanded into a high thin flange; 2) propodeum with posterior transverse carina centrally horizontal; 3) propodeum with lateral longitudinal carina present, defining area dentipara; and 4) ovipositor sheath about times the length of the hind tibia. Distribution. Brazil, Ecuador. New material examined. Brazil: Female. (INPA) Dept. of Amazonas, Borba, Rio Abacaxis, IV Notes. The species is recorded for the first time outside Ecuador. The Brazilian female is similar to the Ecuadorean specimen. FIGURE 4. Propodeum, lateral view, of the amazonica species group (female holotypes). (4) Xanthopimpla amazonica; (b) X. guianensis; (c) X. jussilai; (d) X. pucallpensis; (e) X. vidali. Xanthopimpla pucallpensis Gómez & Sääksjärvi sp. n. (Figs. 2d, 3d, 4d) Type material. Holotype female (BMNH). Peru, Dept. of Ucayali, Pucallpa. J.M. Schuncke leg Female (Fig. 2d). Head in dorsal view moderately short, with genae evenly narrowed behind eyes; frons weakly biconcave; posterior ocellus separate from eye by about 0.8 times its diameter; occipital carina complete, ventrally, laterally and dorsally expanded into a very high thin flange (Fig. 1b); clypeus relatively flat, basally not clearly separated from face; clypeal apex truncate; malar space 0.2 times as long as basal mandibular width; face polished and pubescent, about 1.0 times as wide as medially high. Pronotum with apical edge strongly reflexed and THE NEOTROPICAL SPECIES OF XANTHOPIMPLA Zootaxa 3774 (1) 2014 Magnolia Press 63

8 raised, overlapping propleura. Mesoscutum pubescent, with notauli strongly marked anteriorly, bounded in front by high triangular crests; scutellum convex, with high lateral carinae. Mesopleuron polished, with ventral part densely pubescent; epicnemial carina reaching well above level of lower corner of pronotum, ventrally strongly raised. Metapleuron weakly convex, smooth; submetapleural carina sharp but low, extending back to insertion of hind coxa. Propodeum in profile declivous (Fig. 4d); posterior transverse carina complete and slightly curved; lateromedian longitudinal carina present defining area superomedia (Fig. 3d); lateral longitudinal carina present only anteriorly and posteriorly; pleural carina complete and connected to spiracle by a short carina. Fore wing length about 8 mm; areolet complete; vein Rs slightly sinuous; cu-a opposite Rs&M. Tergite 1 of metasoma about 1.3 times as long as posteriorly broad, with lateral longitudinal and lateromedian longitudinal carina strongly developed anteriorly; tergite 2 about 0.7 times as long as posteriorly broad, with a more or less rhombic raised central area. Claws of hind leg large, without a basal lobe and with four strong hairs in the base. Ovipositor sheath about 0.7 times as long as hind tibia; apex of ovipositor evenly tapered and decurved. Coloration. Pale yellow with the following areas blackish: transverse band across mesoscutum centrally, scuto-scutellar groove, anterior part of propodeum, interocellar area, stripes leading from posterior ocelli to dorsolateral small patches on occipital area and patch on mesosternum. Tergites orange with posterior margins of tergites 1 5 yellowish. Scape yellow in frontal view and brown dorsally. Ovipositor sheath proximally brown, infuscate distally. Wings slightly yellowish, pterostigma yellowish. Male. Unknown Diagnosis. This species can be distinguished from all other Neotropical Xanthopimpla by the combination of the following characters: 1) occipital carina complete and expanded into a high thin flange; 2) propodeum with posterior transverse carina centrally horizontal; and 3) propodeum with lateral carinae not present medially (not defining area dentipara). Xanthopimpla pucallpensis sp. n. resembles X. jussilai; however, the latter has wider tergites and a wider basal area of the propodeum. In addition, the mesosternum of X. jussilai is completely black whereas in X. pucallpensis sp. n. there is only a small black mark. Biological notes. Nothing is known about the hosts of the species. Etymology. The specific name refers to the city of Pucallpa, Department of Ucayali, Peru. Distribution. Peru. Xanthopimpla vidali Gómez sp. n. (Figs. 2e, 3e, 4e) Type material. Holotype female (UNSM): Peru, Dept. Loreto, Reserva Nacional Allpahuayo Mishana, S, W, 124 m, I. Gómez & Sääksjärvi leg. Malaise trap, 28.XI 4.XII Paratypes: one female (UNSM): as above but Malaise trap, XI One female (ZMUT): as above but Malaise trap, 31.X 6.XI Female (Fig. 2e). Head in dorsal view moderately short, with genae evenly narrowed behind eyes; frons biconcave; posterior ocellus separated from eye by 1.0 times its diameter; occipital carina complete, ventrally, laterally and dorsally expanded into a very high thin flange (Fig. 1b); clypeus relatively flat, basally not clearly separated from face; clypeal apex truncate; malar space about 0.3 times as long as basal mandibular width; face polished and pubescent about 1.0 times as wide as medially high. Pronotum with apical edge reflexed and raised, overlapping propleura. Mesoscutum pubescent, with notauli strongly impressed anteriorly, bounded in front by high triangular crests; scutellum convex, with high lateral carinae. Mesopleuron polished, with ventral part densely pubescent; epicnemial carina reaching to well above level of lower corner of pronotum, ventrally strongly raised. Metapleuron weakly convex, smooth; submetapleural carina sharp but low, extending back to insertion of hind coxa. Propodeum in profile strongly declivous (Fig. 4e); posterior transverse carina complete and horizontal; lateromedian longitudinal carina weakly present anteriorly and not present centrally, not defining area superomedia (Fig. 3e); lateral longitudinal carina present, stronger above spiracle; pleural carina complete and connected to spiracle by a short carina. Fore wing length about 10mm; areolet complete; vein Rs sinuous; cu-a opposite or slightly basal to Rs&M; discosubmarginal cell evenly, quite closely hirsute. Tergite 1 of metasoma about 1.3 times as long as posteriorly broad, with lateromedian longitudinal carina strongly developed anteriorly, anterior part of tergite 1 with strong glymma; tergite 2 about 0.6 times as long as posteriorly broad with a more or less rhombic central area. Claws of 64 Zootaxa 3774 (1) 2014 Magnolia Press GÓMEZ ET AL.

9 hind leg large, without a basal lobe and with four strong hairs at the base. Ovipositor sheath about times as long as hind tibia; apex of ovipositor strongly decurved (Fig. 2e), with denticles on upper and lower valve. Coloration. Pale yellow with the following areas blackish: transverse band across mesoscutum centrally, scuto-scutellar groove, anterior part of propodeum, interocellar area, dorso-lateral patches on occipital area, anterior and posterior margins of mesopleuron. Mesosternum yellowish. Tergites orange with posterior margins of tergites 1 6 yellowish. Scape yellow in frontal view and brown dorsally. Antennal flagellum brown, paler ventrally but darker apically, except apical flagellomere distally orange. Ovipositor sheath proximally yellowish, infuscate distally. Wings slightly yellowish, pterostigma yellowish. Male. Unknown. Diagnosis. Xanthopimpla vidali sp. n. can be distinguished from all other Neotropical species of the genus by the combination of the following characters: 1) occipital carina complete and expanded into a high thin flange; 2) propodeum with posterior transverse carina centrally horizontal; 3) ovipositor sheath long, about times the length of the hind tibia; 4) ovipositor tip strongly decurved; and 5) mesosternum yellowish. Biological notes. Nothing is known about the hosts of this species. This species has only been found in the Allpahuayo-Mishana National Reserve. Etymology. This species is named after Vidal Gómez Loarte, father of the first author. Distribution. Peru. The aurita species-group (Figs. 1a, 1d, 1g) Diagnosis. The aurita species-group can be distinguished by the combination of the following characters: 1) occipital carina flange-like only basally (Fig. 1a); 2) propodeal spiracle not joining the pleural carina by a short carina (Fig. 1d); 3) first tergite with lateromedian carinae anteriorly not highly elevated (Fig. 1g); and 4) triangular crests at anterior edge of mesoscutum highly elevated. Remarks. The species of the group are widely distributed in the Neotropical region (Fig. 8) and some are abundant in rainforest habitats. The group was revised previously by Townes (1969). Xanthopimpla allpahuaya Gómez & Sääksjärvi sp. n. (Figs. 5a, 6a, 7a) Type material. Holotype female (UNSM): Peru, Dept. of Loreto, Reserva Nacional Allpahuayo-Mishana S, W, 124 m. Gómez & Sääksjärvi leg. Malaise trap XI Paratypes: One female (UNSM): as above VI One female (UNSM): Peru, Dept. of Loreto, Reserva Nacional Allpahuayo-Mishana. Varillal, Sääksjärvi et al. leg. Malaise trap APHI B3/ XII One female (UNSM): as above but Malaise trap I1(15) 8.XI One male (UNSM): as above but Malaise trap H1(7) 11.VI One male (UNSM): as above but Malaise trap APHI B2/ IX One female (UNSM): as above but Malaise trap I1(12) 19.IX (One female UNSM): as above but Malaise trap H1(10) 2.VIII One female (ZMUT): as above but Malaise trap J1(1) 8.VII One female (ZMUT): as above but Malaise trap D 1/ One female (BMNH): as above Malaise trap I1(11) 18.VII One female (ZMUT): Ecuador, Dept. of Orellana, Onkogane Gare, S, W, 216 m. Terry Erwin leg. Canopy fogging, 29.VI One female (ZMUT): as above but 15.X One female (ZMUT): as above but 6.VII One female (ZMUT): as above but 25.VI One female (INPA): Brazil, Dept. of Amazonas, Manaus. PDBFF, S, W, Klein leg. Malaise trap, 8.X One female (INPA): as above but 19.III One female (INPA): as above but 3.IX One female (INPA): as above but VIII One male (INPA): as above but, 5.XI Female. (Fig. 5a). Head in dorsal view moderately short, with genae evenly narrowed behind eyes; frons weakly biconcave; posterior ocellus separated from eye by 0.7 times its diameter; occipital carina broadly incomplete dorsally, ventrally expanded into a flange; clypeus relatively flat, basally not clearly separated from face, but with a weak transverse groove below level of anterior tentorial pits defining a flat distal portion of the clypeus; clypeal apex truncate; malar space about 0.3 times as long as basal mandibular width; face with THE NEOTROPICAL SPECIES OF XANTHOPIMPLA Zootaxa 3774 (1) 2014 Magnolia Press 65

10 contiguous, coarse punctures. Pronotum with apical edge reflexed and raised. Mesoscutum polished, sparsely punctate, with notauli strongly impressed anteriorly, and bounded in front by a high triangular crests. Scutellum convex, with lateral carinae. Epicnemial carina reaching to above level of lower corner of pronotum. Metapleuron weakly convex, smooth; submetapleural carina sharp but low, extending back to the insertion of hind coxa. Propodeum in profile abruptly declivous; posterior transverse carina complete and curved; lateromedian longitudinal carina only present anteriorly so area superomedia is not defined laterally (Fig. 6a); lateral carina present except above spiracle; pleural carina complete, not connected to spiracle by a short carina. Fore wing length about 10 mm; Rs sinuous; cu-a opposite base of Rs&M; discosubmarginal cell evenly quite closely hirsute. Metasoma with tergite 1 about times as long as posteriorly broad, with well-developed lateral carina anteriorly; tergite 2 about 0.6 times as long as posteriorly broad with a more or less rhombic raised central area which is polished and sparsely punctate laterally. Ovipositor sheath times as long as hind tibia; apex of ovipositor evenly tapered, slightly decurved with denticles on both valves. Coloration. Yellow with the following areas blackish: transverse band centrally across mesoscutum, scutoscutellar groove, anterior part of propodeum; interocellar area, triangular marks on back of head, posterior part of tegula. Anterior and posterior margins of mesopleuron narrowly infuscate; hind coxa ventrally black marked; ovipositor sheath proximally yellowish, strongly infuscate distally. Wings slightly yellowish, pterostigma brownish. Male: Similar to female. Diagnosis. This species can be distinguished from all other Neotropical species of Xanthopimpla by the combination of the following characters: 1) metasoma and hind leg without black marks; 2) propodeum with anterior transverse carina absent centrally; 3) lateromedian longitudinal carinae only present anteriorly (area superomedia not defined laterally); 4) ovipositor sheath about times as long as hind tibia, yellowish but apically infuscate; and 5) occipital carina dorsally absent. Xanthopimpla allpahuaya sp. n. resembles X. aurita especially in propodeal carination. However, this new species has a longer tergite 1, has no black marks on the metasoma and the ovipositor sheath is about half the length of that of X. aurita. Biological notes. This species appears to be restricted to highly vulnerable white sand rain forests (varillal forests). Etymology. The specific name refers to the Allpahuayo-Mishana National Reserve, Department of Loreto, Peru, the principal Peruvian fieldwork area of the first and second authors. This species is named in honour of this unique area of Peruvian Amazonia. Distribution. Brazil, Ecuador, Peru. FIGURE 5. Habitus of the aurita species-group (a) Xanthopimpla allpahuaya; (b) X. aurita; (c) X. craspedoptera; (e) X. rhabdomera; (f) X. spiloptera and the unplaced species (d) X. peruana. 66 Zootaxa 3774 (1) 2014 Magnolia Press GÓMEZ ET AL.

11 FIGURE 6. Propodeum, dorsal view, of the aurita species-group (a) Xanthopimpla allpahuaya; (b) X. aurita; (c) X. craspedoptera; (e) X. rhabdomera; (f) X. spiloptera; and the unplaced species (d) X. peruana. FIGURE 7. Propodeum, lateral view, of the aurita species-group. (a) Xanthopimpla allpahuaya; (b) X. aurita; (c) X. craspedoptera; (e) X. rhabdomera; (f) X. spiloptera; and the unplaced species (d) X. peruana. Xanthopimpla aurita Krieger 1914 (Figs. 5b, 6b, 7b) Original description. See Krieger (1914). Re-descriptions in English. See Townes (1969) and Gauld (1991). Diagnosis. This species can be distinguished from all other Neotropical Xanthopimpla by the combination of the following characters: 1) occipital carina incomplete, expanded into a high thin flange only basally; 2) posterior THE NEOTROPICAL SPECIES OF XANTHOPIMPLA Zootaxa 3774 (1) 2014 Magnolia Press 67

12 transverse carina bowed in the center; 3) ovipositor with 11 teeth on lower valve; and 4) ovipositor sheath about 0.9 to 1.2 times as long as hind tibia. Distribution. Bolivia, Brazil, Colombia, Costa Rica, Ecuador, El Salvador, Mexico, Panama, Peru, Venezuela. Material examined. Brazil: Female (INPA): Dept. of Amazonas, Manaus. Ribeiro, Vidal & Vidal leg. Malaise trap, VII Female (INPA): Dept. of Rondônia, Ouro Preto do Oeste. Rafael & Vidal leg. Malaise trap, VII Female (BMNH): Papagaio. Smith leg Female (BMNH): Dos Arroyos. Smith leg Female (MZUSP): Dept. of São Paulo, Ubatuba. 200 m. Santos & Scott-Santos leg. Malaise trap, I Female (MZUSP): Dept. of São Paulo, Barueri. XII K. Lenko leg. Colombia: Female (BMNH): Putumayo, Villa Garzon. 400 m. Cooper leg Costa Rica: Female (BMNH) Guanacaste National Park, Estacion pitilla, 680 m. Gauld & Mitchell leg. V VI.X Female (BMNH) Guanacaste National Park, Santa Rosa sector, coastal forest m. Gauld & Mitchell leg. VII VIII Female (BMNH) San Antonio de Ecazu, 1300 m. Eberhard & Gauld leg. V Male (BMNH) San Antonio de Ecazu, 1300 m. Eberhard & Gauld leg. XII.1988 I Male (BMNH), Santa Rosa National Park, 0 10 m. Eberhard & Gauld leg. VII Ecuador: Female (BMNH): Sucumbios, Pañacocha. 400 m. Cooper leg Female (NHRS): Napo, Yasuni National Park. Pape & Viklund leg. Malaise trap S, W, 3 20.XI Peru: Female (UNSM): Peru, Dept. of Loreto, Reserva Nacional Allpahuayo-Mishana S, W, 124 m. Gómez & Sääksjärvi leg Malaise trap. XII Female (ZMUT): as above but 28.XI 4.XII Female (ZMUT): as above but 4.X 7.XI Male (ZMUT): as above but 31.X 6.XI Male (ZMUT): as above but 28.XI 4.XII Female (ZMUT): Peru, Dept. of Loreto, Reserva Nacional Allpahuayo-Mishana. Varillal, Sääksjärvi et al. leg Malaise trap E1(8) 29.VI Female (ZMUT): as above but Malaise trap 16.VIII 1.IX Female (ZMUT): as above but Malaise trap I1 20.I Female (ZMUT): as above but Malaise trap F1(15) 8.X Female (ZMUT): as above but Malaise trap G3(16) 21.XII Female (ZMUT): as above but Malaise trap I1(16) 15.XII Female (ZMUT): as above but Malaise trap E2(1) 22.II Female (ZMUT): as above but Malaise trap E1 (8) 29.VI Female (ZMUT): as above but Malaise trap G1 23.I Female (ZMUT): as above but Malaise trap C2(2) 4 18.IX Male (ZMUT): as above but Malaise trap H1.XII Male (ZMUT): as above but Malaise trap E1(17) 28.XII Male (ZMUT): as above but Malaise trap G2(15) 8.XI Male (ZMUT): as above but Malaise trap F1(3) 8.III Male (ZMUT): as above but Malaise trap E1(3) 24.III Male (ZMUT): as above but Malaise trap E1(14) 15.X Male (ZMUT): as above but Malaise trap E1(2) 7.III Male (ZMUT): as above but Malaise trap I Male (ZMUT): as above but Malaise trap E1(11) 18.VIII Male (ZMUT): as above but Malaise trap Female (ZMUT): Peru, Dept. of Madre de Dios, Los Amigos, Isrrael Gómez leg, Malaise trap.vii Notes. The species is recorded here for the first time from Ecuador and Colombia. Xanthopimpla craspedoptera Krieger 1914 (Figs. 5c, 6c, 7c) Original description. See Krieger (1914). Re-descriptions in English. See Townes (1969) and Gauld (1991). Diagnosis. This species can be distinguished from all other Neotropical Xanthopimpla by the combination of the following characters: 1) occipital carina complete, not expanded into a high thin flange; 2) submetapleural carina with a small flange anteriorly; 3) with black spots close to propodeal spiracle and ventrally on hind coxa; and 4) ovipositor sheath about 0.3 to 0.5 times as long as hind tibia. Distribution. Brazil, Colombia, Costa Rica, Ecuador, Panama, Peru. New material examined. Brazil: Female (INPA): Dept. of Amazonas, Iranduba. Vidal leg. Malaise trap, VII Female (INPA): Dept. of Amazonas, Iranduba. Oliveira, Tonon, Somavilla & Azevedo leg. Malaise trap, XI Ecuador: Female (NHRS): Napo, Yasuni National Park. Pape & Viklund leg. Malaise trap S, W, 3 20.XI Male (NHRS): Napo, Yasuni National Park. Pape & Viklund leg. Malaise trap S, W, 3 20.XI Peru: Female (ZMUT): Dept. of Loreto, Reserva Nacional Allpahuayo-Mishana. Varillal, Sääksjärvi et al. leg 68 Zootaxa 3774 (1) 2014 Magnolia Press GÓMEZ ET AL.

13 Malaise trap B2(4) X Female (ZMUT): Dept. of Cusco, Amarakaeri, Castillo leg, Hand net. XI Female (ZMUT): Dept. of Madre de Dios, Los Amigos, Isrrael Gómez leg, Malaise trap.vi Notes. The species is widely distributed in Central and South America but is represented only by a few specimens in collections. Xanthopimpla rhabdomera Townes 1969 (Figs. 5e, 6e, 7e) Original description. See Townes (1969). Diagnosis. This species can be distinguished from all other Neotropical Xanthopimpla by the combination of the following characters: 1) occipital carina complete dorsally; 2) propodeum with anterior transverse carina weak, sometimes discontinuous when joining lateromedian longitudinal carina; and 3) ovipositor sheath uniformly blackish about 0.4 to 0.5 times as long as hind tibia. Distribution. Brazil, Peru. New material examined. Brazil: Female (BMNH): Dept. of Santa Catarina, Nova Teutônia. Plaumann leg Female (BMNH): as above but Peru: Female (ZMUT) Dept. of Junin, Chanchamayo. Isrrael Gómez leg. 806 m. I Notes. The species is recorded for the first time outside of Brazil. Xanthopimpla spiloptera Krieger 1914 (Figs. 5f, 6f, 7f) Original description. See Krieger (1914). Re-description in English. See Townes (1969). Diagnosis. This species can be distinguished from all other Neotropical Xanthopimpla by the combination of the following characters: 1) occipital carina complete dorsally; 2) propodeum with posterior transverse carina strongly angulate; 3) ovipositor sheath about 0.55 to 0.75 times as long as hind tibia; and 4) fore wing strongly infumate apically. Distribution. Brazil, French Guiana, Peru. New material examined. Brazil: Female (BMNH): Dept. of Santa Catarina, Nova Teutônia. Plaumann leg. XI Female (BMNH): as above but 30.VII Female (BMNH): as above but 12.VIII Female (BMNH): as above but VI Female (BMNH): as above but 10.VII Female (BMNH): as above but 12.VI Female (BMNH): as above but 27.IX Female (BMNH): as above but Female (BMNH): as above but French Guiana: Female (ZMUT) Montage des Chevaux. Seag leg. VI Peru: Female (BMNH) Dept. of Junin, coordillera azul J.M. Schunke leg. 760 m. 24.V B.M Notes. The species is recorded from the first time outside of Brazil. Unplaced species (Figs. 1c, 1f, 1i) Xanthopimpla peruana Krieger 1914 (Figs. 5d, 6d, 7d) Original description. See Krieger (1914). Re-description in English. See Townes (1969). Diagnosis. Xanthopimpla peruana can be distinguished from all other Neotropical species by the combination of the following characters: 1) occipital carina complete, rather weak dorsally; 2) propodeum with reduced carination, with only posterior transverse carina present (Fig. 1f); 3) triangular crests at the anterior edge of the THE NEOTROPICAL SPECIES OF XANTHOPIMPLA Zootaxa 3774 (1) 2014 Magnolia Press 69

14 mesoscutum small (Fig. 1c); 4) ventrolateral carina complete across the length of the first tergite (Fig. 1i); and 5) body almost entirely lemon yellowish. Remarks. The Andean species X. peruana is established as an unplaced species outside of the aurita and amazonica species-groups. It may be possible to place it into a new species-group in the future after the tropical Andes become better sampled. Distribution. Ecuador, Peru. New material examined. Peru: Female (UNSM): Dept. of Cusco, Cosñipata, S, W, 1500 m. Castillo leg, Hand net. X Female (ZMUT): as above but IX Male (ZMUT): as above but IX Male (ZMUT): as above but IX Notes. The species seems to be the only Neotropical Xanthopimpla restricted to the tropical Andes at about m above sea level. Discussion The study of Neotropical Xanthopimpla was initiated by Krieger (1914) when he described the first species from the region. Later, Townes (1969) revised the Neotropical Xanthopimpla and described a new species from Brazil, X. rhabdomera. From the limited material available Townes placed all the Neotropical Xanthopimpla species known to him in a single group, the endemic aurita species-group. Recently, Gómez et al. (2009) suggested that there may be more species-groups in the Neotropical region. In the present study, after an examination of all the described and previously undescribed Neotropical species, we define a new species-group and establish one unplaced species which potentially will form a new species-group in the future. FIGURE 8. Distribution of the Xanthopimpla species-groups and X. peruana in the Neotropical region. 70 Zootaxa 3774 (1) 2014 Magnolia Press GÓMEZ ET AL.

15 Distribution and species-richness. In the Neotropical region, the genus Xanthopimpla is currently represented by 11 species belonging to two species-groups and one unplaced species. The amazonica speciesgroup seems to be endemic to northern tropical South America, especially Amazonia. To date, the species of this group have only been found in lowland areas. The aurita species-group is widely distributed across the whole Neotropical region; most of the species have extensive ranges and they have been found at elevations ranging from 0 to about 1600 m. The unplaced X. peruana seems to be restricted to the tropical Andes at elevations ranging from 1200 to about 1600 m. The distribution of the species in the Neotropical region is illustrated in Fig. 8. All the Neotropical Xanthopimpla are present in South America and only two species, X. aurita and X. craspedoptera, are also found in Central America. It appears that the Neotropical Xanthopimpla reach their peak species-richness in Peru with nine of the 11 described species occurring there. However, it has to be taken into account that during the last decade the Peruvian fauna of Ichneumonidae has been studied more intensively in comparison with many other Neotropical countries (e.g. Sääksjärvi et al. 2004; Gómez et al. 2009). Most probably, Xanthopimpla species are wide-spread in the Neotropical region and further sampling in other countries may yield a similar species-richness as found in Peru. Xanthopimpla is primarily a tropical lowland genus. All described Neotropical and Old World species have been found at elevations no higher than 1800 m. In the Neotropical region only X. peruana seems to inhabit exclusively mid altitudes above 1200 m. The vast lowland Amazonian region seems to provide favorable conditions for the diversification of Xanthopimpla in South America. Nevertheless, the number of Xanthopimpla species described from the Neotropical region is only about five per cent of those described from tropical Asia which seems to reflect the Old World biogeographical origin of the genus. Xanthopimpla is the most species-rich pimpline genus. Recently, Pham et al. (2011) described 14 new species from Vietnam. Currently, with the four new species described here, the total number of described species is 265. Most probably the final number of Xanthopimpla species will be much higher as the ichneumonid fauna of most tropical areas (especially the Amazon basin and Congo basin) have only been sampled superficially. The status of the widely distributed Xanthopimpla aurita. Xanthopimpla aurita is the predominant Neotropical species of the genus. Its geographical range is broad and extends from northern Mexico to southern Brazil (Gauld 1991), and from sea level up to about 1300 m. This species is abundant in our Peruvian Amazonian Malaise trap samples (we have collected more than 120 specimens belonging to this species). In addition, we examined specimens from Brazil, Colombia, Costa Rica and Ecuador (deposited in BMHN, INPA, MZUSP and NHRS). In this material, most of the South American specimens agree well with the re-description by Townes (1969), but differ to some degree from that made by Gauld (1991) who examined material mainly from Costa Rica. In Gauld s re-description the specimens are described as having all tergites with black bands (Fig. 5b) whereas in most of the South American specimens only the first tergite possesses a black band. In addition, the ovipositor seems to be longer in South American specimens (ovipositor sheath hind tibia ratio being in South America and in Costa Rica). Gauld (1991) also commented that the material examined by him was limited and because of this it was hard to formulate clear conclusions about the intraspecific variation of this species. There are a few Brazilian specimens with similar metasomal coloration to the Costa Rican specimens. There is also some variation in the fore wing color, with a more yellowish tinge in some Amazonian specimens. Despite this variation we regard all the specimens as belonging to X. aurita confirming the wide distributional range of the species proposed by Gauld (1991). The varying coloration of the species might depend on e.g. different mimicry complexes in the Central and South American parts of its range. However, further morphological and molecular studies are clearly needed to resolve the status of this species in the Neotropical region. Acknowledgements This work was made possible by the Kone Foundation, Finland (under the project 'Biodiversity and multiple trophic interactions' lead by IES). Nhi Thi Pham provided valuable information on Southeast Asian Xanthopimpla. Márcio Oliveira, Terry Erwin, Alexey Reshchikov, Hege Vårdal, Helena Onody, Anu Veijalainen, Gerardo Lamas and Yves Braet (SEAG team) facilitated Neotropical material. Andrew Bennet provided photos of paratypes deposited in the Canadian National Collection of Insects (CNC). We are grateful to Edgard Palacio and John Jennings (University of Adelaide, South Australia) for valuable comments on the manuscript. The Ministry of THE NEOTROPICAL SPECIES OF XANTHOPIMPLA Zootaxa 3774 (1) 2014 Magnolia Press 71