Zootaxa 3337: 1–56 (2012) ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Article ZOOTAXA Copyright © 2012 · Magnolia Press ISSN 1175-5334 (online edition)

Revision of the Neotropical species of Trichacis Foerster (: Platygastroidea: ), with description of 24 new species

TANIA MILENA ARIAS-PENNA1, LUBOMIR MASNER2 & THIBAUT DELSINNE3 1. Universidad Nacional de Colombia, Bogotá D.C., COLOMBIA. E-mail: [email protected] 2. Agriculture and Agri-Food Canada, Ottawa, Ontario K1A OC6, CANADA. E-mail: [email protected] 3. Biological Evaluation Section, Royal Belgian Institute of Natural Sciences, 29 rue Vautier, 1000 Brussels, BELGIUM. E-mail: [email protected]

Table of contents

Abstract ...... 1 Introduction ...... 2 Material and methods ...... 3 Trichacis ...... 3 Key to Neotropical species of Trichacis ...... 4 Trichacis acarinata Arias-Penna & Masner, new species...... 5 Trichacis acuminata Arias-Penna & Masner, new species ...... 7 Trichacis acuta Arias-Penna & Masner, new species ...... 8 Trichacis ariaspennae Buhl 2011 ...... 9 Trichacis clypeata Arias-Penna & Masner, new species...... 10 Trichacis colombiana Arias-Penna & Masner, new species ...... 11 Trichacis concavata Arias-Penna & Masner, new species ...... 14 Trichacis corrugata Arias-Penna & Masner, new species ...... 14 Trichacis costaricana Arias-Penna & Masner, new species ...... 15 Trichacis delsinnei Arias-Penna & Masner, new species ...... 17 Trichacis depressa Arias-Penna & Masner, new species ...... 17 Trichacis dianae Arias-Penna & Masner, new species ...... 19 Trichacis fernandezi Arias-Penna & Masner, new species ...... 21 Trichacis hansoni Arias-Penna & Masner, new species ...... 21 Trichacis kaulbarsi Arias-Penna & Masner, new species ...... 23 Trichacis laticornis Buhl 2001 ...... 26 Trichacis magnifica Arias-Penna & Masner, new species ...... 26 Trichacis meridionalis (Brues) 1910 ...... 28 Trichacis mexicana Arias-Penna & Masner, new species ...... 28 Trichacis panamana Arias-Penna & Masner, new species ...... 30 Trichacis pecki Arias-Penna & Masner, new species ...... 32 Trichacis procera Arias-Penna & Masner, new species ...... 32 Trichacis proximata Arias-Penna & Masner, new species ...... 34 Trichacis punctata Arias-Penna & Masner, new species ...... 34 Trichacis sculpturata Arias-Penna & Masner, new species ...... 37 Trichacis transversata Arias-Penna & Masner, new species ...... 39 Trichacis triangulata Arias-Penna & Masner, new species ...... 39 Acknowledgements ...... 43 References ...... 43

Abstract

The hymenopteran genus Trichacis is presumed to be endoparasitoid of Cecidomyiidae. Only three species of this genus were hitherto recognized in the Neotropical region. Here, twenty four (24) new species are described based on 145 specimens: T. acarinata (Costa Rica), T. acuminata (Bolivia), T. acuta (Colombia), T. clypeata (Costa Rica), T.

Accepted by I. Miko: 28 Mar. 2012; published: 7 Jun. 2012 1 colombiana (Colombia), T. concavata (Costa Rica), T. corrugata (El Salvador, Mexico), T. costaricana (Costa Rica), T. delsinnei (Costa Rica), T. depressa (Costa Rica), T. dianae (Colombia, Ecuador, Venezuela), T. fernandezi (Ecuador), T. hansoni (Brazil, Costa Rica, Nicaragua, Panama), T. kaulbarsi (Mexico), T. magnifica (Mexico), T. mexicana (Mexico), T. panamana (Panama), T. pecki (Ecuador), T. procera (Mexico), T. proximata (Costa Rica), T. punctata (Brazil), T. sculpturata (Mexico), T. transversata (Costa Rica), and T. triangulata (Mexico). A key for males and females of the 27 Neotropical species is provided.

Key words: Central America, identification key, parasitoid, South America,

Introduction

All members of the hymenopteran genus Trichacis Foerster (1856) are presumed to be koinobiont endoparasitoids of gall midge larvae (Diptera: Cecidomyiidae) (Masner 1983; Masner & Hanson 2006). Because most of the latter feed on living plants and can be important agricultural pests (Gagné & Jaschhof 2009), Trichacis species have the potential to be key agents in biological control (Masner 1983). However, the biology and diversity of Trichacis are poorly known, especially in Central and South America, where only three species were recognized before the present study: Trichacis meridionalis (Brues 1910), T. laticornis Buhl 2001, and T. ariaspennae Buhl 2011. The last revision of Trichacis focused on the Nearctic species (Masner 1983). In his work, Masner (1983) followed the classification proposed by Kozlov (1970), and placed Trichacis in the tribe Platygastrini of the subfamily , based on the palpal and tibial spur formulae (2–1, and 1–2–2, respectively). He recognized 15 species, described the characters delimiting the genus, and discussed those allowing to distinguish Trichacis from three morphologically close Platygastrinae genera, Isocybus Foerster, Metanopedias Brues and Trichacoides Dodd. These characters are the presence of a specialized area filled with a tuft of fine dense erect hairs on the top of the mesoscutellum, a smooth frons, receding temples behing the eyes, a 3–5 segmented antennal clava in females, and six visible tergites in female metasoma. In the most recently published phylogeny of the Platygastroidea (Murphy et al. 2007), Trichacis formed a clade with Synopeas Foerster (Platygastridae: Platygastrinae), and Trichacis + Synopeas with Piestopleura Foerster (Platygastridae: Platygastrinae). Isocybus, Metanopedias and Trichacoides were not included in the analysis. In this paper, we described 24 new species. Specimens were collected from Mexico to Bolivia and Brazil, and from 200 to 2600 meters in elevation. The type material of Trichacis meridionalis, T. laticornis and T. ariaspennae was not examined but we included these species in our work based on the original descriptions and Dr. Buhl's personal communication.

Material and methods

Material. This work is based on the examination of 145 specimens (111 females and 34 males) from the Canadian National Collection of , Arachnids and Nematodes, Ottawa, Canada (CNCI). All specimens were dried and pinned. Holotypes and paratypes were deposited at the CNCI, at the Natural History Museum of London, United Kingdom (BMNH), and at the Instituto de Ciencias Naturales–Museo de Historia Natural, Universidad Nacional de Colombia, Bogotá D.C., Colombia (ICN).

Observations and images. Observations, measurements (Appendix 1) were made at 160x on a Leica S8APO stereomicroscope. Bright field images were taken using a Leica DFC290 camera attached to a Leica Z6 APO stereomicroscope. Series of images were taken by focusing the sharpness on different levels of the structure, using the Leica Application Suite v38 (2003–2011), and combined with the “Align and balance used frame (quick)” and “Do stack” commands of CombineZP (Hadley 2010). Final processing of images was done in Adobe Photoshop CS. Original images are deposited in Morphbank (collection number: 791583; http://www.morphbank.net/ 791583).

2 · Zootaxa 3337 © 2012 Magnolia Press ARIAS-PENNA ET AL. Abbreviations.

Collecting methods are abbreviated as follows: FIT—flight interception trap, MT—malaise trap, SS—screen sweeping, and YPT—yellow pan trap. Terminology and abbreviations of morphological characters follow Masner (1983), Masner and Huggert (1989) and morphological terms used in this revision were matched to the Hymenoptera Anatomy Ontology (HAO, Yoder et al. 2010, http://api.hymao.org/projects/32/public/ontology/) (Appendix 2). Surface sculpture terminology follows Harris (1979).

A—Antennomeres: sclerites of the antenna; they are numbered from the scape (A1) to the apical most sclerite (A10). EH—Eye height: in frontal view, the distance from the lowermost to the uppermost point of the eye. INA—Inter-notaular area: the area on the mesoscutum delimited laterally by the notauli. IOS—Interorbital space: in frontal view, the shortest distance between the inner margins of the eyes. LNA—Lateral notaular area: the area on the mesoscutum delimited medially by the notauli. LOL—Lateral ocellar line: the shortest distance on the vertex between the inner margins of the anterior ocellus and the posterior ocelli. OOL—Ocular ocellar line: the shortest distance between the posterior ocellus and the eye. POL—Posterior ocellar line: the shortest distance between the posterior ocelli. T—Tergite (T1, T2, T3…): the dorsal sclerite of the metasomal segment.

Results

Trichacis Förster Trichacis: Förster, 1856: 108, 115. Original description. Type: Platygaster pisis Walker, designated by Ashmead (1893). Keyed. Thomson, 1859: 70. Description, spelling error, keyed; Marshall, 1873: 20. Catalog of species of Britain; Walker, 1873: 540, 541. Keyed; Howard, 1886: 174. Keyed; Cresson, 1887: 85, 249. Keyed, catalog of species of U.S. and Canada; Provancher, 1888: 403. Description; Ashmead, 1893: 263, 264, 294. Description, keyed, key to species of U.S. and Canada; Marchal, 1897: 86. Description; Dalla Torre, 1898: 481. Catalog of species; Ashmead, 1900: 330. List of species of West Indies, spelling error; Ashmead, 1903: 98, 99. Keyed; Kieffer, 1914: 361. Keyed; Brues, 1916: 531. Keyed; Kieffer, 1916: 564. Description; Fouts, 1924: 3, 13. Description, key to species of U.S. and Canada, keyed; Kieffer, 1926: 563, 710. Description, keyed, key to species; Morley, 1929: 47. Catalog of species of Britain; Jansson, 1939: 175. Keyed; Maneval, 1940: 117. Keyed; Mani, 1941: 31, 33. Catalog of species of India, keyed; Muesebeck & Walkley, 1951: 709. Catalog of species of U.S. and Canada; Muesebeck & Walkley, 1956: 404. Citation of type species; Muesebeck & Masner, 1967: 302. Second supplement to Muesebeck & Walkley (1951); Kozlov, 1971: 59. Keyed. Vlug, 1973: 183. Types of Platygastridae in Vienna; Fergusson, 1978: 121. Kozlov, 1978: 664. Keyed; Checklist of species of Britain; Muesebeck, 1979: 1176. Catalog of species of America North of Mexico; De Santis, 1980: 322. Catalog of species of Brazil; Mani & Sharma, 1982: 204, 215. Description, keyed; Masner, 1983: 1071. Diagnosis, description, key to species of the Nearctic region; Vlug, 1985: 214. Key to types of species of Walker; Kozlov, 1995: 126. Keyed; Vlug, 1995: 84. Catalog of world species; Austin & Field, 1997: 63, 68. Structure of ovipositor system, discussion of phylogenetic relationships; Buhl, 1999: 41. Key to species of Fennoscandia and Denmark; Buhl, 2001: 34, 35. Description; Loiácono & Margaría, 2002: 556. Catalog of Brazilian species; Arias-Penna, 2007: 11. Neotropical distribution, new distribution record for Bolivia, Ecuador, Mexico, Panama, and Venezuela; Buhl 2011: 121, 122. Description.

(Taxonomic synopsis modified from http://osuc.biosci.ohio-state.edu/hymDB/ eol_scelionidae.content_page?page_level=3&page_id=taxon_page_data&page_version=7852)

NEOTROPICAL TRICHACIS Zootaxa 3337 © 2012 Magnolia Press · 3 Diagnosis The presence of the specialized area on the top of the mesoscutellum allows separation of Trichacis from Piestopleura and Synopeas, as well as from all other platygastrines (Masner 1983). In addition, Synopeas can be distinguished from Trichacis by its median keels of propodeum contiguous (wide apart in Trichacis), its short mesoscutellar projection never with tuft of hairs, and the fusion of its first and second metasomal tergites. Piestopleura differs from Trichacis by the extreme lateral compression of its thorax, its anteroposteriorly compressed head and the dorsoventrally compressed metasoma (Jackson 1969).

Comments As noted for the Nearctic species (Masner 1983), Neotropical members of Trichacis form a morphologically homogeneous complex with few diagnostic characters. In this study, we confirm the usefulness of the characters listed by Masner (1983) for species separation, such as the presence/absence of temple projection and the sculpture of the anteromedial area of the second metasomal tergite. In addition, we recognized the following characters as important: presence/absence of the hyperoccipital carina (Figs 21A, 21B); the extent of the hyperoccipital carina (Figs 21B–21E); type of sculpture on the vertex; extent of sculpture on the vertex; distance separating the posterior ocelli from the hyperoccipital carina; extent of the sculpture on the inter-notaular area; shape of the specialized area of the mesoscutellum; shape of the median projection of the mesoscutellum. However, we did not find consistent trends in the variation of pilosity, and we did not treat this character in our descriptions. Similarly, every specimen included in the present study had a trifid foretibial spur, shallow micropunctuation along the hind margin of the second metasomal tergite, and three long setae below the tegula. Trichacis, like other Platygastroidea (Masner 1993), is sexually dimorphic. Males can be distinguished easily from females by the filiform antenna, by the enlarged antennomere 4 (= flagellomere 2), and by the externally visible metasomal tergite 7. One of the possible synapomorphies shared by all members of Platygastroidea is the presence of basiconic sensilla on each clavomere of the female antenna (Masner 1993). However, in all the Neotropical Trichacis females we examined, the sensilla were entirely sunken into the antenna and therefore not visible. This is also found in two Nearctic species: Trichacis celticola Masner and T. elongata Masner (Masner 1983).

Key to Neotropical Species of Trichacis

1 Vertex and occiput smooth ...... 2 - Vertex and/or occiput at least partially coriaceous or coriaceous with striae ...... 4 2 (1) Hyperoccipital carina absent (Fig. 21A), Costa Rica ...... Trichacis acarinata n. sp. ♀ - Hyperoccipital carina present ...... 3 3 (2) OOL longer than LOL, T1 shorter than wide, Costa Rica ...... Trichacis proximata n. sp. ♀ - OOL slightly shorter than LOL, T1 as wide as long, Brazil ...... Trichacis meridionalis (Brues) ♀ 4 (1) Hyperoccipital carina absent (Figs 17C, 17D); Brazil ...... Trichacis punctata n. sp. ♀ - Hyperoccipital carina present (Figs 21B–21E) ...... 5 5 (4) Hyperoccipital carina incomplete or discontinuous, but not reaching eye margin (Figs 21C–21E) ...... 6 - Hyperoccipital carina complete, reaching eye margin (Fig. 21B) ...... 22 6 (5) In dorsal view, hyperoccipital carina fading before or until reaching level of imaginary line connecting eye inner margin to posterior cephalic margin (Fig. 21C) ...... 7 - In dorsal view, hyperoccipital carina surpassing level of imaginary line connecting eye inner margin to posterior cephalic margin (Figs 21D, 21E)...... 11 7 (6) Temple projection present (Figs 2B, 3A, 3B) ...... 8 - Temple projection absent ...... 9 8 (7) Area of lateral vertex between posterior ocellus and hyperoccipital carina coriaceous (Fig. 2B); and between hyperoccipital carina and occipital carina coriaceous with longitudinal striae (Fig. 1H); Bolivia ...... Trichacis acuminata n. sp. ♀♂ - Lateral vertex coriaceous, without striae; Colombia...... Trichacis acuta n. sp. ♀ 9 (7) Inter-notaular area entirely coriaceous (Fig. 17G); Mexico ...... Trichacis sculpturata n. sp. ♀ - Inter-notaular area with only anterior 1/3 coriaceous, other 2/3 smooth ...... 10 10 (9) Combined length of T3–T6 less than length of T2 (Figs 12A, 12E); specialized area of mesoscutellum with cone-like elevation (Fig. 12B); metasoma not depressed dorsoventrally; Mexico ...... Trichacis magnifica n. sp. ♀♂ - Combined length of T3–T6 greater than length of T2 (Fig. 7F); specialized area of mesoscutellum heart-shaped, moderately elevated; metasoma depressed dorsoventrally (Fig. 8A); Costa Rica...... Trichacis depressa n. sp. ♀ 11 (6) Mesopleural carina complete (Fig. 9A); vertex laterally coriaceous, and with well-marked longitudinal striae between hyperoccipital carina and occipital carina (Fig. 8D); Colombia, Ecuador, Venezuela ...... Trichacis dianae n. sp. ♀♂ - Mesopleural carina incomplete (Fig. 18E) ...... 12 12 (11) Hyperoccipital carina curving up and merging with stria running along the eye inner margin (Fig. 21E) ...... 13

4 · Zootaxa 3337 © 2012 Magnolia Press ARIAS-PENNA ET AL. - Hyperoccipital carina not curving up and not merging with other stria (Fig. 21D) ...... 15 13 (12) Anteromedial area of T2 (i.e. area situated between anterior pits) slightly concave and with longitudinal striae (Fig. 9F); Ecuador ...... Trichacis fernandezi n. sp. ♀ - T2 anteromedially (between pits) flattened and smooth ...... 14 14 (13) Lateral region of vertex between posterior ocellus and hyperoccipital carina weakly coriaceous, and with cluster of 4-6 well- marked longitudinal curved striae, which seem to emerge from hyperoccipital carina; lateral vertex area between hyperoccipital carina and occipital carina coriaceous (Fig. 15A); Ecuador ...... Trichacis pecki n. sp. ♀ - Lateral region of vertex between posterior ocellus and hyperoccipital carina markedly coriaceous, with 9-12 well-separated and irregularly curved longitudinal striae, and 2-3 curved transverse striae running between outer margin of posterior ocellus and inner margin of eye; lateral vertex area between hyperoccipital carina and occipital carina coriaceous, and with well- marked longitudinal striae (Fig. 7C); Costa Rica ...... Trichacis delsinnei n. sp. ♀ 15 (12) Posteromedial region of inter-notaular area broadly concave (Figs 5E, 5G); Costa Rica ...... Trichacis concavata n. sp. ♀ - Posteromedial region of inter-notaular area convex or blunt ...... 16 16 (14) Inter-notaular area entirely coriaceous (Figs 6A, 6D); El Salvador; Mexico ...... Trichacis corrugata n. sp. ♀ - Inter-notaular area anterior 1/3 coriaceous, otherwise smooth ...... 17 17 (16) T1 shorter than wide (Figs 13F, 14B) ...... 18 - T1 at least as long as wide...... 19 18 (17) Faint, transverse, slightly curved striae present throughout the anterior region of vertex; distance between posterior ocellus and hyperoccipital carina longer than posterior ocellus diameter (Fig. 14D); Panama...... Trichacis panamana n. sp. ♀♂ - Presence of faint, transverse, slightly curved striae restricted to interocellar area; distance between posterior ocellus and hyperoccipital carina shorter than posterior ocellus diameter (Fig. 13C); Mexico ...... Trichacis mexicana n. sp. ♀♂ 19 (17) Length and width of T1 equal (Fig. 5A); all lateral vertex region coriaceous ...... 20 - T1 longer than wide (Fig.18C); lateral region of vertex betweeen posterior ocellus and hyperoccipital carina coriaceous, and between hyperoccipital carina and occipital carina with well-marked longitudinal striae (Fig. 18E); Costa Rica ...... Trichacis transversata n. sp. ♀ 20 (19) Notaulus present (Fig. 9H) ...... 21 - Notaulus absent (Fig. 5A); Colombia ...... Trichacis colombiana n. sp. ♀ 21 (20) Vertex laterovertically striate-coriaceous; Honduras ...... T. laticornis Buhl 2001 - Vertex laterally coriaceous, without striae; Brazil, Costa Rica, Nicaragua, Panama ...... Trichacis hansoni n. sp. ♀ ♂ 22 (5) Mesopleural carina complete (Fig. 15F); Mexico ...... Trichacis procera n. sp. ♀ - Mesopleural carina incomplete (Fig. 20E) ...... 23 23 (22) Notaulus absent (Fig. 11A); Mexico ...... Trichacis kaulbarsi n. sp. ♀♂ - Notaulus present ...... 24 24 (23) Clypeus fused with interantennal process, forming rounded yellow-brown projected sheet (Fig. 4B); Costa Rica ...... Trichacis clypeata n. sp. ♀♂ - Clypeus and interantennal process well-separated, not forming projecting lamella ...... 25 25 (24) Combined length of T3–T6 less than length of T2 ...... 26 - Combined length of T3–T6 greater than length of T2; Brazil ...... Trichacis ariaspennae Buhl 2011 26 (25) Distance between posterior ocellus and hyperoccipital carina less than diameter of posterior ocellus (Fig. 20C); specialized area of mesoscutellum triangular and moderately elevated (Figs 18H; 20E); Mexico ...... Trichacis triangulata n. sp. ♀♂ - Distance between posterior ocellus and hyperoccipital carina greater than diameter of posterior ocellus; specialized area of mesoscutellum circular and elevated (Fig. 6F); Costa Rica ...... Trichacis costaricana n. sp. ♀

Descriptions of Neotropical Trichacis species

Trichacis acarinata Arias-Penna & Masner, new species Figures 1A–1F, 21A

Female. Body length: 2.80 mm (holotype). Dark brown; A1–A6 yellow; A7–A10 brown; mandible yellow with base and apex brown; leg yellow; hind coxa brown; fore wing infuscate. Head, in dorsal view, 2.2 times as wide as long; EH < IOS; interocellar area smooth; OOL > LOL; antennal clava 3-segmented; A1 not reaching vertex dorsal margin; A2–A5, A10 longer than wide; A6–A9 wider than long; base of mandible smooth; frons above torulus with transverse striae; interantennal process surpassing ventral border of torulus; clypeus exposed; clypeal margin concave medially; gena and vertex smooth; temple projection absent; hyperoccipital carina absent; distance between posterior ocellus and occipital carina line > OOL. Mesosoma slightly arched; antero-admedian line absent; notaulus anteriorly incomplete; mesoscutum smooth; posteromedial region of inter-notaular area acute; parapsidal line present; central pronotal area smooth; specialized area of mesoscutellum heart-shaped; median projection of specialized area of mesoscutellum triangular; mesopleural carina incomplete (length of mesopleural carina/length of mesopleuron = 0.42); fore wing not surpassing apex of metasoma.

NEOTROPICAL TRICHACIS Zootaxa 3337 © 2012 Magnolia Press · 5 FIGURE 1. Bright field images of Trichacis species. A–F: Trichacis acarinata Arias-Penna & Masner, n. sp., ♀ (holotype). A, head, frontal view; B, habitus, dorsal view; C, habitus, lateral view; D, head and prothorax, anterolateral view; E head and mesosoma, lateral view; F, head, mesosoma and T1, dorsal view. G, H: Trichacis acuminata Arias-Penna & Masner, n. sp., ♀ (holotype). G, head and mesosoma, anterior view; H, habitus, dorsal view.

6 · Zootaxa 3337 © 2012 Magnolia Press ARIAS-PENNA ET AL. Metasoma distinctly elongate; combined length of T3–T6 > T2; T1 wider than long; T2, T5 and T6 longer than wide; T3–T4 wider than long; T2 anteromedial area (i.e. between anterior pits) smooth and concave; T2–T3 smooth; anterior half of T4 with shallow punctuation; anterior half of T5 and T6 densely and deeply punctulate. Male: Unknown. Etymology. The name refers to the absence of hyperoccipital carina. Diagnosis. Trichacis acarinata and T. punctata are characterized by the absence of a hyperoccipital carina. Trichacis acarinata differs from T. punctata by the following characters: combined length of T3–T6 > T2; mesoscutum smooth; posteromedial region of inter-notaular area acute. Material examined. Holotype: 1 ♀, COSTA RICA, Punt. San Vito, Biol. Sta. Las Alturas, 1500 m, May 1992, MT, P. Hanson (CNCI). Holotype is deposited in CNCI.

Trichacis acuminata Arias-Penna & Masner, new species Figures 1G, 1H; 2A–2C

Female. Body length: 1.80 mm (holotype); 2.25 mm (paratype); mean length ± SD = 2.03 ± 0.32 mm. Black; radicle and base of A1 yellow; distal part of A1 brown and A2–A10 dark brown; mandible brown with apex dark brown; fore leg yellow except apical tarsi brown; mid leg brown except basal 2/3 of tibia and 1–4 tarsi yellow; hind leg brown, base of trochanter, apical 2/3 of femur, apical 2/3 of tibia and tarsi darker; coxae dark brown; fore wing infuscate in posterior 2/3, with darker and lighter patterns in basal 1/3. Head, in dorsal view, two times as wide as long; EH < IOS; interocellar area finely coriaceous; OOL > LOL; antennal clava 4-segmented; A1 not reaching vertex dorsal margin; A2–A5 longer than wide; A6 as long as wide; A7–A9 wider than long; base of mandible smooth; frons above torulus coriaceous, and with transverse striae; interantennal process smooth, slightly curved down and surpassing ventral border of torulus; clypeus exposed; clypeal margin truncate; gena smooth; lateral parts of vertex coriaceous between posterior ocellus and hyperoccipital carina, and coriaceous and longitudinally striate between hyperoccipital carina and occipital carina; temple projection present; hyperoccipital carina incomplete, fading until reaching level of imaginary line connecting eye inner margin to posterior cephalic margin; distance between posterior ocellus and hyperoccipital carina longer than posterior ocellus diameter. Mesosoma slightly arched; antero-admedian line present; notaulus anteriorly incomplete; lateral notaular area with coriaceous sculpture along its anterior margin and inter-notaular area with anterior 1/3 coriaceous and posterior 2/3 smooth; posteromedial region of inter-notaular area convex; parapsidal line present; central pronotal area smooth; specialized area of mesoscutellum heart-shaped and elevated; median projection of specialized area of mesoscutellum subtriangular; mesopleural carina incomplete (length of mesopleural carina/length of mesopleuron = 0.52); fore wing distinctly surpassing apex of metasoma. Metasoma elongate; combined length of T3–T6 < T2; T1 as long as wide; T2 longer than wide; T3–T6 wider than long; anteromedial part of T2 smooth and concave; T6 subtriangular; T3–T6 densely and deeply punctulate. Male. Body length: 1.90 mm (allotype). Similar to female except the following: A1–A5 yellow; A6–A10 brown; fore and mid legs yellow; hind leg yellow except brown coxa, tarsal segments and apical 1/3 of femur and tibia; wing infuscate in posterior 2/3, with darker and lighter patterns in basal 1/3; antenna filiform; A2–A4, A6–A10 longer than wide; A5 as long as wide. Etymology. The name of the species refers to the temple projection. Diagnosis. Both Trichacis acuminata and T. acuta have temple projection but may be distinguished by the sculpture on the medial area of vertex, which is covered by striae in T. acuminata and smooth in T. acuta, and by the shape of the anteromedial part of T2, which is concave in T. acuminata and flat in T. acuta. Interestingly, the Neartic species Trichacis bison, T. cornuta, and T. dracula also have temple projection (Masner 1983). Material examined. Holotype: 1 ♀; BOLIVIA, Cochabamba, montane & cloud forest, 17º08´52´´S 65º42´54´´W, 1800 m, February 6–8 1999, FIT, F. Génier (CNCI); allotype: 1 ♂, La Paz, Chulumani, Apa-Apa, 16º22´S 67º30´W, 1800 m, May 1–4 1997, L. Masner (CNCI); paratype: 1 ♀, with same data as holotype (ICN). Holotype is deposited in CNCI.

NEOTROPICAL TRICHACIS Zootaxa 3337 © 2012 Magnolia Press · 7 FIGURE 2. Bright field images of Trichacis species. A–C: Trichacis acuminata Arias-Penna & Masner, n. sp. A, ♀, habitus, lateral view (holotype); B, ♀, head and anterior mesosoma, dorsal view (holotype); C, ♂, head and mesosoma, anteroventral view (allotype). D–F: Trichacis acuta Arias-Penna & Masner, n. sp., ♀ (holotype). D, head, anterior view; E, habitus, dorsal view; F, habitus, lateral view.

Trichacis acuta Arias-Penna & Masner, new species Figures 2D–2F; 3A–3D; 21C

Female. Body length: 2.50 mm (holotype); 1.80–2.40 mm (paratypes, n=7); mean length ± SD = 2.18 ± 0.24 mm. Black; base of A1 yellow, distal part of A1 and A2–A10 dark brown; mandible yellow but apex brown and base dark brown; leg dark brown; fore wing infuscate in posterior 2/3, with darker and lighter patterns in basal 1/3.

8 · Zootaxa 3337 © 2012 Magnolia Press ARIAS-PENNA ET AL. Head, in dorsal view, two times as wide as long; EH < IOS; interocellar area coriaceous; OOL > LOL; antennal clava 4-segmented; A1 not reaching vertex dorsal margin; A2–A6, A10 longer than wide; A7 as long as wide; A8–A9 wider than long; base of mandible smooth; frons above torulus with 7–8 weak transverse striae; interantennal process smooth and reaching, but not surpassing, ventral border of torulus; clypeus exposed; clypeal margin convex medially; gena striate; vertex laterally coriaceous and medially smooth; temple projection present; hyperoccipital carina incomplete: in dorsal view, hyperoccipital carina fading until reaching level of imaginary line connecting eye inner margin to posterior cephalic margin; distance between posterior ocellus and hyperoccipital carina longer than posterior ocellus diameter. Mesosoma arched; antero-admedian line present; notaulus anteriorly incomplete; lateral notaular area with coriaceous sculpture along its anterior margin; inter-notaular area with anterior 1/3 coriaceous, otherwise smooth; posteromedial region of inter-notaular area convex; parapsidal line present; central pronotal area smooth; specialized area of mesoscutellum heart-shaped; median projection of specialized area of mesoscutellum subtriangular; mesopleural carina incomplete (length of mesopleural carina/length of mesopleuron = 0.55); fore wing distinctly surpassing apex of metasoma. Metasoma elongate; combined length of T3–T6 < T2; T1 as wide as long, T2 longer than wide, anteromedial area of T2 smooth and flattened; T3–T6 shallowly covered by dense punctuation; T6 triangular. Male. Body length: 1.90 mm (allotype); 1.80 mm (paratypes, n=2); mean length ± SD = 1.83 ± 0.06 mm. Similar to female except the following: antenna filiform; A1–A10 dark brown; A1 distinctly surpassing vertex margin; A5 as long as wide; A8–A9 longer than wide; T1 shorter than wide; T2 longer than wide; T7 shallowly covered by dense punctuation; T6 wider than long. Etymology. The name of the species refers to the temple projection. Diagnosis. Both Trichacis acuminata and T. acuta have temple projection but sculpture on the medial area of vertex is covered by striae in T. acuminata and smooth in T. acuta, and the shape of the anteromedial part of T2 is concave in T. acuminata and flat in T. acuta. The Neartic species Trichacis bison, T. cornuta, and T. dracula also have temple projection (Masner 1983). Material examined. Holotype: 1 ♀, COLOMBIA, Chuza, June 18 1992, MT, E. Palacio (CNCI); allotype: 1 ♂ with same data as holotype but caught July 27 1992 (CNCI); paratypes: 5 ♀♀, with same data as allotype (4 at CNCI and 1 at ICN); 2 ♀♀, with same data as holotype (CNCI); 1 ♂ with same data as holotype (ICN); 1 ♂, with same data as allotype (CNCI). Holotype is deposited in CNCI.

Trichacis ariaspennae Buhl 2011

Trichacis ariaspennae Buhl 2011: 121, 122. Original description.

Diagnosis. Occiput medially smooth, laterally dull reticulate-coriaceous with vertical elements; hyperoccipital carina distinct and complete; OOL:LOL = 3:3.5; distance between posterior ocellus and hyperoccipital carina longer than diameter of posterior ocellus; notaulus strong, ending abruptly very shortly before reaching anterior margin; mesoscutum with very few setae, smooth except for reticulation in anterior 0.2; mid lobe posteriorly narrowly pointed, slightly prolonged to base of mesoscutellum; combined length of T3–T6 > T2 (Buhl 2011 and pers. comm.). The female holotype (the only specimen currently known) was collected in Brazil and was deposited at BMNH (Buhl 2011). Trichacis ariaspennae has a complete hyperoccipital carina, but differs from other species with this character because: T kaulbarsi n sp. has the notaulus absent; T clypeata n. sp. has a projected clypeus fused with interantennal process; T. proximata n sp. has posterior ocellus very close to, almost touching hyperoccipital carina; T. triangulata n. sp. and T. costaricencis n. sp. have combined length of T3–T6 < T2.

NEOTROPICAL TRICHACIS Zootaxa 3337 © 2012 Magnolia Press · 9 FIGURE 3. Bright field images of Trichacis species. A–D: Trichacis acuta Arias-Penna & Masner, n. sp. A, ♀, head and mesosoma anterior view (holotype); B, ♀, head and anterior mesosoma, dorsal view (holotype); C, ♂, head, anterolateral view (allotype); D, ♂, posterior thorax and metasoma, dorsal view (allotype). E, F: Trichacis clypeata Arias-Penna & Masner, n. sp., ♀ (holotype). E, head, anterior view; F, habitus, dorsal view.

Trichacis clypeata Arias-Penna & Masner, new species Figures 3E, 3F; 4A–4E

Female. Body length: 2.50 mm (holotype); 1.90–2.70 mm (paratypes, n=4); mean length ± SD = 2.32 ± 0.30 mm. Black; base of A1, A2–A5 yellow and distal part of A1, A6–A10 brown; mandible yellow with apex brown; fore

10 · Zootaxa 3337 © 2012 Magnolia Press ARIAS-PENNA ET AL. and mid legs yellow; hind leg mostly yellow, femur and tibia apices brown; fore and mid coxae brown; hind coxa dark brown; fore wing infuscate in posterior 2/3, with darker and lighter patterns in basal 1/3. Head, in dorsal view, 2.5 times as wide as long; EH < IOS; interocellar area smooth; OOL > LOL; antennal clava 3-segmented; A1 not reaching vertex dorsal margin; A2–A5, A9, A10 longer than wide; A6–A8 wider than long; base of mandible smooth; frons above torulus with transverse striae; clypeus fused with interantennal process, forming rounded yellow-brown projected sheet; clypeal margin convex medially; gena striate; lateral region of vertex between hyperoccipital carina and occipital carina coriaceous; temple projection absent; hyperoccipital carina complete (i.e. reaching inner margin of eye); posterior ocellus almost reaching hyperoccipital carina. Mesosoma arched; antero-admedian line present; notaulus percurrent; lateral notaular area smooth; inter- notaular area with posterior 2/3 smooth and with coriaceous sculpture in basal 1/3; shape of posteromedial region of inter-notaular area V-like but with blunt tip; parapsidal line present; central pronotal area smooth; specialized area of mesoscutellum heart-shaped and elevated; median projection of specialized area of mesoscutellum broadly triangular; mesopleural carina incomplete (length of mesopleural carina/length of mesopleuron = 0.44); fore wing distinctly surpassing apex of metasoma. Metasoma distinctly elongate; combined length of T3–T6 < T2; T1 as long as wide; T2–T3 longer than wide; T4–T6 shorter than wide; anteromedial area of T2 smooth and flattened; T3 densely covered by shallow punctuation but smooth along its anterior margin, T4–T6 densely covered by shallow punctuation; T6 subtriangular. Male. Body length: 2 mm (allotype). Similar to female except the following: base of A1 yellow; distal part of A1 and A2–A10 brown; fore and mid legs yellow; hind leg brown; coxae brown; wing infuscate in posterior 2/3, with darker and lighter patterns in basal 1/3; antenna filiform; A1 not reaching vertex dorsal margin; A2, A4, A5, A8, A10 longer than wide; A3, A6, A7, A9 as long as wide; T1 almost as long as wide; T2 longer than wide; T3–T7 wider than long; T2 anteromedial area smooth and flattened; T3 densely covered by shallow punctuation but smooth along its anterior margin, T4–T7 densely covered by deep punctuation. Etymology. The name refers to the clypeus, which is fused with the interantennal process in this species. Diagnosis. Trichacis clypeata is the only species that presents the clypeus fused with interantennal process. Material examined. Holotype: 1 ♀, COSTA RICA, Alajuela, Est. Biol. San Ramón, 400 m, October–December 1995, P. Hanson (CNCI); allotype: 1 ♂, Limón, 16 km W Guapiles, 400 m, December 1989, MT, P. Hanson (CNCI); paratypes: 2 ♀♀, with same data as holotype but collected at 900 m, August–September 1995 (CNCI), 1 ♀, with same data as holotype (CNCI); 1 ♀, with same data as holotype but collected at 900 m, July–August 1995 (ICN). Holotype is deposited in CNCI.

Trichacis colombiana Arias-Penna & Masner, new species Figures 4F, 5A–5C

Female. Body length: 1.95 mm (holotype). Black; antenna base of A1 yellow, distal part of A1 brown and A2–A10 dark brown; apex and base of mandible brown, rest yellow; fore leg yellow; mid leg with trochanter, femur and apical 2/3 of tibia brown, otherwise yellow; hind leg dark brown; fore coxa brown; mid and hind coxae dark brown; fore wing infuscate. Head, in dorsal view, 1.7 times as wide as long; EH < IOS; interocellar area finely coriaceous; OOL > LOL; antennal clava 4-segmented; A1 not reaching vertex dorsal margin; A2–A6, A10 longer than wide; A7–A8 wider than long; A9 as long as wide; base of mandible smooth; frons above torulus with 3 transverse striae and coriaceous sculpture; interantennal process smooth and reaching, but no surpassing, ventral border of torulus; clypeus exposed; clypeal margin truncate; gena striate; vertex laterally coriaceous and medially smooth; temple projection absent; hyperoccipital carina incomplete: in dorsal view, hyperoccipital carina surpassing level of imaginary line connecting eye inner margin to posterior cephalic margin; distance between posterior ocellus and hyperoccipital carina longer than posterior ocellus diameter.

NEOTROPICAL TRICHACIS Zootaxa 3337 © 2012 Magnolia Press · 11 FIGURE 4. Bright field images of Trichacis species. A–E: Trichacis clypeata Arias-Penna & Masner, n. sp. A, ♀, habitus, lateral view (holotype); B, ♀, head and anterior mesosoma, lateral view (holotype); C, ♀, head and anterior mesosoma, dorsal view (holotype); D, ♂, head, anterior view (allotype); E, ♂, posterior mesosoma and metasoma, dorsal view (allotype). F: Trichacis colombiana Arias-Penna & Masner, n. sp., ♀ (holotype), head and ventral mesosoma, anterior view.

Mesosoma arched; antero-admedian line present; notaulus absent; posteromedial region of inter-notaular area convex, with anterior region coriaceous and rest smooth; parapsidal line present; central pronotal area smooth; specialized area of mesoscutellum heart-shaped; median projection of specialized area of mesoscutellum triangular; mesopleural carina incomplete; fore wing distinctly surpassing apex of metasoma. Metasoma moderately elongate; combined length of T3–T6 < T2; T1 as long as wide; T2 longer than wide; anteromedial region of T2 smooth and flattened; T3 smooth along its anterior margin and rest densely covered by shallow punctuation; T4–T6 densely covered by deep punctuation; T6 triangular.

12 · Zootaxa 3337 © 2012 Magnolia Press ARIAS-PENNA ET AL. FIGURE 5. Bright field images of Trichacis species. A–C: Trichacis colombiana Arias-Penna & Masner, n. sp., ♀ (holotype). A, habitus, dorsal view; B, habitus, lateral view; C, head and anterior mesosoma, anterodorsal view. D–G: Trichacis concavata Arias- Penna & Masner, n. sp., ♀ (holotype). D, head and ventral mesosoma, anterior view; E, habitus, dorsal view; F, habitus, lateral view; G, head and mesosoma, dorsal view. H: Trichacis corrugata Arias-Penna & Masner, n. sp., ♀ (holotype), head, anterior view.

NEOTROPICAL TRICHACIS Zootaxa 3337 © 2012 Magnolia Press · 13 Male. Unknown. Etymology. Named after the country of the type locality. Diagnosis. Trichacis colombiana and T. kaulbarsi have notaulus absent, but contrary to T. kaulbarsi, T. colombiana has area between anterior and posterior ocelli finely coriaceous and incomplete hyperoccipital carina. Material examined. Holotype: 1 ♀, COLOMBIA, Magdalena, PNN Santa Marta, El Ramo, 2500 m, 10°4´N 73°35´W, July 15–31 2000, MT, J. Cantillo (CNCI). Holotype is deposited in CNCI.

Trichacis concavata Arias-Penna & Masner, new species Figures 5D–5G

Female. Body length: 1.40 mm (holotype). Black shining; A1–A6 yellow; A7–A10 brown; mandible black with apex brown; fore and mid legs yellow and hind leg brown; coxae dark brown; fore wing infuscate. Head, in dorsal view, 2.2 times wider than long; EH < IOS; interocellar area finely coriaceous; OOL = LOL; antennal clava 3-segmented; A1 not reaching vertex dorsal margin; A2–A4, A8, A10 longer than wide; A5–A6 as long as wide; A7–A9 wider than long; base of mandible smooth; frons above torulus and interantennal process with three marked transverse striae; interantennal process smooth and surpassing ventral border of torulus; clypeus hidden by interantennal process; clypeal margin truncate; gena striate; vertex laterally coriaceous and medially smooth; temple projection absent; hyperoccipital carina incomplete: in dorsal view, hyperoccipital carina surpassing level of imaginary line connecting eye inner margin to posterior cephalic margin; distance between posterior ocellus and hyperoccipital carina shorter than posterior ocellus diameter. Mesosoma arched; antero-admedian line present; notaulus anteriorly incomplete and weakly marked; lateral notaular area with coriaceous sculpture along its anterior margin; inter-notaular area with coriaceous sculpture in 1/ 3 anterior, and with posterior 2/3 smooth; posteromedial region of inter-notaular area broadly concave; parapsidal line present; central pronotal area smooth; specialized area of mesoscutellum heart-shaped; median projection of specialized area of mesoscutellum subtriangular; mesopleural carina incomplete (length of mesopleural carina/ length of mesopleuron = 0.72); fore wing distinctly surpassing apex of metasoma. Metasoma moderately elongate; combined length of T3–T6 < T2; T1 longer than wide; T2 slightly shorter than wide; T3–T5 wider than long; anteromedial area of T2 smooth and slightly concave; T3 densely covered by shallow punctuation; T4–T6 densely covered by deep punctuation; T6 subtriangular. Male. Unknown. Etymology. The name of the species refers to the concavity of the posteromedial region of the inter-notaular area. Diagnosis. Trichacis concavata is the single species with posteromedial region of inter-notaular area concave, and with OOL = LOL. Material examined. Holotype: 1 ♀, COSTA RICA, Alajuela, Reserva Rincón Forestal, Estación Caribe, 10º53´N 85º18´W, 400 m, February 19–20 2003, SS, J.S. Noyes (CNCI). Holotype is deposited in CNCI.

Trichacis corrugata Arias-Penna & Masner, new species Figures 5H; 6A–6D

Female. Body length: 2.30 mm (holotype); 2.45 mm (paratype); mean length ± SD = 2.38 ± 0.11 mm. Black; A1–A6 yellow; A7–A10 brown; apex and basis of mandible brown, otherwise yellow; fore and mid legs yellow; hind leg with trochanter, femur, and basal 2/3 of tibia yellow, otherwise brown; coxae yellow; fore wing infuscate. Head, in dorsal view, 2.2 times as wide as long; EH < IOS; interocellar area coriaceous; OOL > LOL; antennal clava 4-segmented; A1 not reaching vertex dorsal margin; A2–A6, A10 longer than wide; A7 as long as wide; A8–A9 wider than long; base of mandible smooth; frons above interantennal process and torulus with five deeply

14 · Zootaxa 3337 © 2012 Magnolia Press ARIAS-PENNA ET AL. marked transverse striae, which almost reach inner margin of eye; interantennal process striate and surpassing ventral border of torulus; clypeus exposed; clypeal margin convex medially; gena striate; vertex laterally coriaceous; medial area of vertex covered by transverse striae between posterior ocellus and hyperoccipital carina, and smooth between hyperoccipital carina and occipital carina; temple projection absent; hyperoccipital carina incomplete: in dorsal view, hyperoccipital carina surpassing level of imaginary line connecting eye inner margin to posterior cephalic margin; distance between posterior ocellus and hyperoccipital carina longer than posterior ocellus diameter. Mesosoma arched; antero-admedian line present; notaulus anteriorly incomplete; lateral notaular area with coriaceous sculpture along its anterior margin; inter-notaular area coriaceous; posteromedial region of inter- notaular area convex; parapsidal line present; area of the pronotum between epomiae with anterior region smooth and rest coriaceous; specialized area of mesoscutellum heart-shaped and elevated; median projection of specialized area of mesoscutellum broadly triangular and curving up; mesopleural carina incomplete (length of mesopleural carina/length of mesopleuron = 0.34); fore wing distinctly surpassing apex of metasoma. Metasoma elongate; combined length of T3–T6 < T2; T1 longer than wide; anteromedial area of T2 smooth and concave; T3 smooth along its anterior margin; T4–T6 densely covered by shallow punctuation; T6 subtriangular. Male. Unknown. Etymology. The name of the species refers to the striae on the vertex. Diagnosis. T. corrugata and T. sculpturata have inter-notaular area coriaceous. The shape of the specialized area of mesoscutellum and the sculpture on vertex allow to separate these species. Material examined. Holotype: 1 ♀, MEXICO, Tamps, Río Frío, nr. Gomez Farias, June 6 1983, UV. S. Peck & M. Kaulbars (CNCI); paratype: 1 ♀, EL SALVADOR: Santa Ana, Guachipilin, Montecristo, 1200 m, 14°27’N 89°22’W, August 2000, R. Menjivar, MT (CNCI). Holotype is deposited in CNCI.

Trichacis costaricana Arias-Penna & Masner, new species Figures 6E–6G

Female. Body length: 2.60 mm (holotype). Black; A1–A5 yellow; A6–A10 brown; mandible brown with base and apex dark brown; fore and mid legs yellow; hind leg tarsi, apex of femur and apex of tibia brown, otherwise yellow; fore and mid coxae brown; hind coxa dark brown; fore wing infuscate. Head, in dorsal view, 2.2 times as wide as long; EH < IOS; interocellar area with fine coriaceous sculpture; OOL < LOL; antennal clava 4-segmented; A1 not reaching vertex dorsal margin; A2–A4, A7, A10 longer than wide; A5 as long as wide; A6, A8–A9 wider than long; base of mandible smooth; frons above torulus, with transverse striae and coriaceous; clypeus exposed; clypeal margin convex medially; interantennal process striate, curving down, and surpassing anterior ventral of torulus; gena smooth; lateral region of vertex coriaceous, and with longitudinal striae between hyperoccipital carina and occipital carina; temple projection absent; hyperoccipital carina complete; distance between posterior ocellus and hyperoccipital carina longer than posterior ocellus diameter. Mesosoma slightly arched; antero-admedian line present; notaulus anteriorly incomplete; lateral notaular area with coriaceous sculpture along its anterior margin; basal 1/3 of inter-notaular area coriaceous, rest smooth; posteromedial region of inter-notaular area convex; parapsidal line present; central pronotal area smooth; specialized area of mesoscutellum circular and elevated; mesopleural carina incomplete (length of mesopleural carina/length of mesopleuron = 0.55); fore wing slightly surpassing apex of metasoma. Metasoma elongate; combined length of T3–T6 < T2; T1 shorter than wide; T2 longer than wide; T3 wider than long; anteromedial area of T2 smooth and concave; T3–T6 wider than long; T3 with some parts smooth and densely covered by shallow punctuation; T4–T6 densely covered by deep punctuation; T6 subtriangular. Male. Unknown. Etymology. Named after the country of the type locality.

NEOTROPICAL TRICHACIS Zootaxa 3337 © 2012 Magnolia Press · 15 FIGURE 6. Bright field images of Trichacis species. A–D: Trichacis corrugata Arias-Penna & Masner, n. sp. ♀ (holotype). A, habitus, dorsal view; B, habitus, lateral view; C, head and anterior mesosoma, dorsal view; D, head and mesosoma, dorsolateral view. E–G: Trichacis costaricana Arias-Penna & Masner, n. sp., ♀ (holotype). E, head and anteroventral mesosoma, anterior view; F, habitus, dorsal view; G, habitus, lateral view. H: Trichacis delsinnei Arias-Penna & Masner, n. sp., ♀ (holotype), head, anterior view.

16 · Zootaxa 3337 © 2012 Magnolia Press ARIAS-PENNA ET AL. Diagnosis. Trichacis costaricana, like T. kaulbarsi, has its specialized area of mesoscutellum circular and elevated, but can be distinguished by the following characters: notaulus present; OOL < LOL and distance between posterior ocellus and hyperoccipital carina longer than posterior ocellus diameter. Material examined. Holotype: 1 ♀, COSTA RICA, Guanacaste, N. P. Biol. Sta. Cacao, 1200 m, February 13 1995, L. Masner (CNCI). Holotype is deposited in CNCI.

Trichacis delsinnei Arias-Penna & Masner, new species Figures 6H; 7A–7D

Female. Body length: 2.20 mm (holotype). Black; A1–A6 yellow; A7–A10 brown; mandible brown with base and apex dark brown; fore leg yellow; mid and hind legs with tarsi and tibial apex brown, otherwise yellow; coxae yellow; fore wing infuscate. Head, in dorsal view, 1.9 times as wide as long; EH < IOS; interocellar area with fine coriaceous sculpture; OOL > LOL; antennal clava 3-segmented; A1 not reaching vertex dorsal margin; A2–A5, A10, A6–A9 as long as wide; base of mandible smooth; frons above torulus coriaceous with 4 marked transverse striae, and above interantennal process weakly striate; interantennal process smooth and surpassing ventral border of toruli; clypeus exposed; clypeal margin convex medially; gena smooth; lateral region of vertex between posterior ocellus and hyperoccipital carina markedly coriaceous, with 9-12 well-separated and irregularly curved longitudinal striae, and 2-3 curved transverse striae running between outer margin of posterior ocellus and inner margin of eye; medial area of vertex smooth; temple projection absent; hyperoccipital carina incomplete, but curving up and merging with stria running along the eye inner margin; distance between posterior ocellus and hyperoccipital carina longer than posterior ocellus diameter. Mesosoma slightly arched; antero-admedian line present; notaulus anteriorly incomplete; lateral notaular area with coriaceous sculpture along its anterior margin; inter-notaular area with anterior 1/3 coriaceous and smooth in its posterior 2/3; posteromedial region of inter-notaular area broadly convex; parapsidal line present; area of the pronotum between epomiae smooth; specialized area of mesoscutellum heart-shaped and elevated; mesopleural carina incomplete (length of mesopleural carina/length of mesopleuron = 0.39); fore wing slightly surpassing apex of metasoma. Metasoma elongate; combined length of T3–T6 < T2; T1, T2 shorter than wide; T3–T6 wider than long; anteromedial area of T2 smooth and flattened; T3 finely and densely punctured; T4–T5 and T6 anterior border densely covered by deep punctuation; T6 subtriangular. Male. Unknown. Etymology. The species is named in honor of the entomologist Thibaut Dominique Delsinne. Diagnosis. Trichacis delsinnei, T. pecki and T. fernandezi have an incomplete hyperoccipital carina, which curves up before reaching the eye margin. T. delsinnei is diagnosed by its vertex sculpture and by the broad convexity of its posterior inter-notaular area. Material examined. Holotype: 1 ♀, COSTA RICA, Alajuela, Est. Biol. San Ramón, 900 m, August–November 1995, MT, P. Hanson (CNCI). Holotype is deposited in CNCI.

Trichacis depressa Arias-Penna & Masner, new species Figures 7E, 7F; 8A, 8B

Female. Body length: 2.40 mm (holotype); 1.90–2.70 mm (paratypes, n=4); mean length ± SD = 2.22 ± 0.33 mm. Black; A1–A10 dark brown; mandible yellow with apex brown; fore leg yellow; mid leg with trochanter, femur and base of tibia yellow, otherwise brown; hind leg dark brown; mid coxa yellow; hind coxa brown; fore wing infuscate.

NEOTROPICAL TRICHACIS Zootaxa 3337 © 2012 Magnolia Press · 17 FIGURE 7. Bright field images of Trichacis species. A–D: Trichacis delsinnei Arias-Penna & Masner, n. sp., ♀ (holotype). A, habitus, dorsal view; B, habitus, lateral view; C, head, dorsal view; D, head and mesosoma, dorsal view. E, F: Trichacis depressa Arias-Penna & Masner, n. sp., ♀ (holotype). E, head and fore legs, anterior view; F, habitus, dorsal view.

Head, in dorsal view, 2.1 times wider than long; EH > IOS; interocellar area with fine coriaceous sculpture; OOL > LOL; antennal clava 3-segmented; A1 distinctly surpassing vertex dorsal margin; A2–A10 longer than wide; base of mandible smooth; frons between external margin of torulus and inner margin of eye medially covered by delicate coriaceous sculpture; interantennal process truncate, smooth and reaching, but not surpassing, ventral border of torulus; clypeus exposed; clypeal margin truncate; gena smooth; vertex laterally coriaceous and medially smooth; temple projection absent; hyperoccipital carina incomplete: in dorsal view, hyperoccipital carina fading before reaching level of imaginary line connecting eye inner margin to posterior cephalic margin; distance between posterior ocellus and hyperoccipital carina shorter than posterior ocellus diameter.

18 · Zootaxa 3337 © 2012 Magnolia Press ARIAS-PENNA ET AL. Mesosoma slightly arched; antero-admedian line present; notaulus anteriorly incomplete; lateral notaular area with coriaceous sculpture along its anterior margin; inter-notaular area with anterior 1/3 coriaceous and smooth in its posterior 2/3; posteromedial region of inter-notaular area broadly convex; parapsidal line present; central pronotal area smooth; specialized area of mesoscutellum heart-shaped, moderately elevated, with a rather broad tuft of hairs; mesopleural carina incomplete (length of mesopleural carina/length of mesopleuron = 0.12); fore wing not surpassing apex of metasoma. Metasoma distinctly elongate and depressed dorsoventrally; combined length of T3–T6 > T2; T1–T2 longer than wide; T3–T6 wider than long; anteromedial area of T2 smooth and flattened; T3–T5 with some parts smooth and densely covered by shallow punctuation; T6 densely covered by shallow punctuation along its anterior margin and rest smooth; T6 subtriangular. Male. Unknown. Etymology. The name of the species refers to its depressed metasoma. Diagnosis. Trichacis depressa and T. acarinata have combined length of T3–T6 > T2. T. depressa is hitherto the only Neotropical species with a depressed metasoma and with EH > IOS. Material examined. Holotype: 1 ♀, COSTA RICA, San José, Cerro de la Muerte, 19 km S, 3 km W Empalme, 2600 m, April–July 1992, MT, P. Hanson (CNCI); paratypes: 3 ♀♀, with same data as holotype, (2 at CNCI and 1 at ICN); 1 ♀ S.J. Cerro de la Muerte. November 1992, 2600 m, MT, P. Hanson (CNCI). Holotype is deposited in CNCI.

Trichacis dianae Arias-Penna & Masner, new species Figures 8C–8F; 9A

Female. Body length: 1.90 mm (holotype); 1.50–2 mm (paratypes, n=20); mean length ± SD = 1.83 ± 0.13 mm. Black; A1–A5 yellow; A6–A10 brown; mandible yellow with apex brown; fore and mid legs yellow; hind leg brown but with apices of femur and tibia darker; coxae dark brown; fore wing infuscate. Head, in dorsal view, 2.7 times as wide as long; EH < IOS; interocellar area with fine coriaceous sculpture; OOL < LOL; antennal clava 4-segmented; A1 not reaching vertex dorsal margin; A2–A3, A5, A10 longer than wide; A4, A6–A7 as long as wide; A8–A9 wider than long; base of mandible smooth; frons above torulus and interantennal process with delicate transverse striae and coriaceous sculpture; interantennal process striate and surpassing ventral border of torulus; clypeus exposed; clypeal margin convex medially; gena striate; lateral area of vertex weakly coriaceous between posterior ocellus and hyperoccipital carina and with well-marked longitudinal striae between hyperoccipital carina and occipital carina; temple projection absent; hyperoccipital carina almost complete: in dorsal view, hyperoccipital carina surpassing level of imaginary line connecting eye inner margin to posterior cephalic margin; distance between posterior ocellus and hyperoccipital carina equal or slightly longer than posterior ocellus diameter. Mesosoma arched; antero-admedian line present; notaulus anteriorly incomplete; lateral notaular area with coriaceous sculpture along its anterior margin; inter-notaular area coriaceous in its basal 1/3 and smooth in its posterior 2/3; posteromedial region of inter-notaular area convex; parapsidial line present; central pronotal area smooth; specialized area of mesoscutellum heart-shaped, median projection of specialized area of mesoscutellum triangular; mesopleural carina complete; fore wing distinctly surpassing apex of metasoma. Metasoma elongate; combined length of T3–T6 < T2; T1 as wide as long; T2 longer than wide; T3–T6 wider than long; anteromedial area of T2 smooth and concave; anterior pit of T2 with longitudinal striae; T3 smooth along its anterior margin and densely covered by shallow punctuation; T4–T6 densely covered by deep punctuation; T6 subtriangular. Male. Body length: 2 mm (allotype); 1.70–1.85 mm (paratypes, n=4); mean length ± SD = 1.82 ± 0.12 mm. Similar to female except the following: antenna filiform and yellow; A2 wider than long; A4, A6–A10 longer than wide; T1 as wide as long; T3–T7 densely covered by deep punctuation. Etymology. The species is named in honor of the entomologist Diana Carolina Arias-Penna. Diagnosis. Trichacis dianae and T. procera have complete mesopleural carina. T. dianae differs by the combination of the following characters: OOL < LOL; presence of longitudinal striae on vertex; anterior pit of T2 with longitudinal striae; smooth and concave T2 anteromedial area.

NEOTROPICAL TRICHACIS Zootaxa 3337 © 2012 Magnolia Press · 19 FIGURE 8. Bright field images of Trichacis species. A, B: Trichacis depressa Arias-Penna & Masner, n. sp., ♀ (holotype). A, habitus, lateral view; B, head and anterior mesosoma, dorsal view. C–F: Trichacis dianae Arias-Penna & Masner, n. sp., ♀ (holotype). C, head, anterior view; D, habitus, dorsal view; E, habitus, lateral view; F, head, dorsal view.

Material examined. Holotype: 1 ♀, VENEZUELA, Lara, Yacambu, N. P., 1200 m, May 10 1981, H. Townes (CNCI); allotype: 1 ♂, with same data as holotype but caught May 7 1981 (CNCI); paratypes: COLOMBIA: 1 ♂, Magdalena, 11º10´N 74º8´W, 800 m, April 26–30 1973, J. Helava (CNCI); ECUADOR: 1 ♀, Sucumbios, Sacha Lodge, 0º30´S 76º30´W, 270 m, August 16–23 1994, MT, P. Hibbs (CNCI); 1 ♀, Sucumbios, Río Napo, Sacha Lodge, 0º30´S 76º30´W, 270 m, July 25–August 3 1994, MT, Peter Hibbs (CNCI); 1 ♀, Napo, Limoncocha, 250 m, June 15–28 1976, S. & J. Peck (CNCI); VENEZUELA: 2 ♂, 3 ♀♀ with the same data as allotype (CNCI); 1 ♂, 6 ♀♀, with same data as holotype (CNCI); 4 ♀, with same data as holotype but caught May 13 1981 (CNCI), 2 ♀♀ Merida, Merida–Sta. Rosa, ca. 1800 m, May 11 1981. L. Masner (CNCI); 1 ♀, Zulia El Tucuco, 200 m, Primary

20 · Zootaxa 3337 © 2012 Magnolia Press ARIAS-PENNA ET AL. rain for., April 1981, L. Masner (CNCI); 1 ♀, Aragua, Parque Nac. H. Pittier, Rancho Grande env., 1100 m, April 10–14 1994, YPT, L. Masner (ICN). Holotype is deposited in CNCI.

Trichacis fernandezi Arias-Penna & Masner, new species Figures 9B–9F; 21E

Female. Body length: 1.60 mm (holotype). Black; A1–A5 yellow; A6–A10 brown; mandible yellow with base and apex brown; fore and mid legs yellow; hind leg with tarsi and apical ½ of femur and tibia brown, otherwise yellow; fore wing infuscate. Head, in dorsal view, two times as wide as long; EH < IOS; interocellar area finely coriaceous; OOL < LOL; antennal clava 4-segmented; A1 not reaching vertex dorsal margin; A2, A3, A10 longer than wide; A4, A6 as long as wide; A5, A7–A9 wider than long; base of mandible striate; frons above torulus with three transverse striae and coriaceous sculpture; interantennal process with reticulate striae, surpassing ventral border of torulus; clypeus hidden by interantennal process; clypeal margin truncate; gena striate; lateral region of vertex, between external margin of posterior ocellus and eye inner margin, coriaceous, with 2–4 curved transverse striae; lateral region of vertex between hyperoccipital carina and occipital carina markedly coriaceous and longitudinally striate; temple projection absent; hyperoccipital carina incomplete but curving up and merging with stria running along the eye inner margin; distance between posterior ocellus and hyperoccipital carina longer than posterior ocellus diameter. Mesosoma slightly arched; antero-admedian line present; notaulus anteriorly incomplete; lateral notaular area with coriaceous sculpture along its anterior margin; inter-notaular area with anterior 1/3 coriaceous and smooth in its posterior 2/3; posteromedial region of inter-notaular area narrow and convex; parapsidal line present; central pronotal area smooth; specialized area of mesoscutellum heart-shaped; median projection of specialized area of mesoscutellum subrectangular; mesopleural carina incomplete (length of mesopleural carina/length of mesopleuron = 0.45); fore wing slightly surpassing apex of metasoma. Metasoma elongate; combined length of T3–T6 < T2; T1 shorter than wide; T2 longer than wide; T3 wider than long; anteromedial area of T2 slightly concave and longitudinally striate; T3 smooth along its anterior margin and rest with shallow punctuation; T4–T6 densely covered by deep punctuation; T6 subtriangular. Male. Unknown. Etymology. The species is named in honor of the entomologist Fernando Fernández. Diagnosis. T. fernandezi and T. pecki have an incomplete hyperoccipital carina of similar shape. Trichacis fernandezi has OOL < LOL, a slightly concave and longitudinally striate anteromedial area of T2, and an inter- notaular area that is narrow and convex posteriorly. Material examined. Holotype: 1 ♀, ECUADOR, Sucumbios, Sacha Lodge, 0º30´S 76º30´W, July 3–13 1994, MT, Peter Hibbs (CNCI). Holotype is deposited in CNCI.

Trichacis hansoni Arias-Penna & Masner, new species Figures 9G, 9H; 10A–10E

Female. Body length: 2.20 mm (holotype); 1.85–2.60 mm (paratypes, n=37); mean length ± SD = 2.22 ± 0.17 mm. Black; A1 yellow; A2–A10 brown; mandible yellow with base and apex brown; leg yellow; fore wing infuscate. Head, in dorsal view, 2.1 times as wide as long; EH < IOS; interocellar area finely coriaceous; OOL > LOL; antennal clava 3-segmented; A1 not reaching vertex dorsal margin; A2–A6, A8, A10 longer than wide; A7 as long as wide; A9 wider than long; base of mandible smooth; frons above interantennal process and torulus with four transverse striae; interantennal process with transverse striae and reaching, but not surpassing, ventral border of torulus; clypeus exposed; clypeal margin weakly convex; gena striate; vertex laterally coriaceous and medially smooth; temple projection absent; hyperoccipital carina incomplete: in dorsal view, hyperoccipital carina surpassing level of imaginary line connecting eye inner margin to posterior cephalic margin; distance between posterior ocellus and hyperoccipital carina shorter than posterior ocellus diameter.

NEOTROPICAL TRICHACIS Zootaxa 3337 © 2012 Magnolia Press · 21 FIGURE 9. Bright field images of Trichacis species. A: Trichacis dianae Arias-Penna & Masner, n. sp., ♂ (Allotype), head, mesosoma and anterior metasoma, lateral view. B–F: Trichacis fernandezi Arias-Penna & Masner, n. sp., ♀ (holotype). B, head and ventral mesosoma, anterior view; C, habitus, dorsal view; D, habitus, lateral view; E, head, dorsal view; F, head, mesosoma and anterior metasoma, dorsal view. G, H: Trichacis hansoni Arias-Penna & Masner, n. sp., ♀ (holotype). G, head, anterior view; H, habitus, dorsal view.

22 · Zootaxa 3337 © 2012 Magnolia Press ARIAS-PENNA ET AL. Mesosoma slightly arched; antero-admedian line weakly marked; notaulus almost percurrent; lateral notaular area with coriaceous sculpture along its anterior margin; inter-notaular area with basal 1/3 coriaceous, and posterior 2/3 smooth; posteromedial region of inter-notaular area convex; parapsidal line present; area of the pronotum between epomiae smooth; specialized area of mesoscutellum heart-shaped, moderately elevated; median projection of specialized area of mesoscutellum rectangular, borded laterally by a broad tuft of hairs; mesopleural carina incomplete (length of mesopleural carina/length of mesopleuron = 0.25); fore wing slightly surpassing apex of metasoma. Metasoma elongate; combined length of T3–T6 < T2; T1, T2 as long as wide; T3–T6 wider than long; T2 anteromedial area smooth and flattened; T3 smooth along its margin anterior and rest densely covered by shallow punctuation; T4–T6 densely covered by deep punctuation; T6 subtriangular. Male. Body length: 2.20 mm (allotype); 2–2.30 mm (paratypes, n=6); mean length ± SD = 2.17 ± 0.10 mm. Similar to female except the following: antenna filiform, A1–A10 brown, fore and mid legs, including coxae, yellow; hind leg with coxa dark brown, trochanter and femur yellow, tibia and tarsi brown; A7–A9 longer than wide. Etymology. Named after Paul E. Hanson, the collector of numerous specimens of this species, including the holotype and the allotype. Diagnosis. Trichacis hansoni is the single species with the combined characters: T1 and T2 as long as wide; specialized area of mesoscutellum heart-shaped, moderately elevated, with a rectangular median projection bordered laterally by a broad tuft of hairs, distance between posterior ocellus and hyperoccipital carina shorter than posterior ocellus diameter. Material examined. Holotype: 1 ♀, COSTA RICA, San José, Zurqui de Moravia, 1600 m, April 1995, MT, P. Hanson (CNCI); allotype: 1 ♂. Punt. San Vito, Biol. Sta Las Alturas, 2050 m, September–November 1992, P. Hanson (CNCI); paratypes: 2 ♀♀, with same data as holotype (CNCI); 1 ♀, with same data as holotype but captured in September 1996 (CNCI); 1 ♀, with same data as holotype but caught in October 1995 (ICN); 9 ♀♀, Heredia, Porrosati, 10º05´34´´N 84º06´41´´W, 1900 m, August 22 1988, YPT, L. Masner (CNCI); 3 ♀♀, Heredia, Río Ceruelas, 10º05´41´´N 84º06´41´´W, 1600 m, August 22–30 1988, YPT, creek, L. Masner (CNCI); 10 ♀♀, with same data as allotype (CNCI); 2 ♀♀, Cartago, La Congrega, 1950 m, June–July 1992, P. Hanson (CNCI); 1 ♀, Cartago, Cangreja, 1950 m, July 1991, MT, P. Hanson (CNCI); 1 ♂, Guan. GCA. Est Pitilla, 700 m, February 12–27 1996, J. S. Noyes. MT/YPT (CNCI); 3 ♀♀, San Vito de C.B., Las Cruces, 1200 m, July 9–August 7 1982 (CNCI); 1 ♀, BRAZIL: Est. Rio de Janeiro, Silva Jardim, August 1974. F.M. Oliveira (CNCI); 1 ♂, NICARAGUA: 50 km E Matagalpa, El Coyolar, 800 m, June 1991, at light, S. Hue (CNCI); 3 ♂♂, 1 ♀, PANAMA: Chiriquí, 6 km NE Boquete, 1650 m, June 1996, FIT, J. Ashe & Brooks (CNCI); 1 ♀, Chiriquí, La Fortuna, Hydro trail, 1450 m, May 23–June 9 1996, FIT, J. Ashe & Brooks (CNCI); 1 ♀, Chiriquí, Prov. 2 km W Cerro Punta, 1700 m, May 19–June 1977, Peck & Howden (CNCI); 1 ♂, Cerro Campana, 825 m, June 1–5 1995, FIT, J. Ashe & Brooks (CNCI); 1 ♀, Chiriquí Cerro Pando, 8°54´N 82°47´W, 1875 m, July 17–18 1996, FIT, J. Ashe & Brooks (CNCI). Holotype is deposited in CNCI.

Trichacis kaulbarsi Arias-Penna & Masner, new species Figures 10F; 11A–11G

Female. Body length: 1.60 mm (holotype). Black; A1–A10 brown and mandible yellow with apex brown; leg yellow; coxa dark brown; fore wing infuscate. Head, in dorsal view, 1.9 times as wide as long; EH < IOS; interocellar area smooth; OOL > LOL; antennal clava 4-segmented; A1 not reaching vertex dorsal margin; A2–A5, A8–A10 longer than wide; A6–A7 as long as wide; base of mandible smooth; frons above interantennal process and torulus with four deeply marked transverse striae; interantennal process smooth and surpassing ventral border of torulus; clypeus exposed; clypeal margin truncate; gena smooth; lateral region of vertex smooth between posterior ocellus and hyperoccipital carina, and with well-marked longitudinal striae between hyperoccipital carina and occipital carina; temple projection absent; hyperoccipital carina complete, distance between posterior ocellus and hyperoccipital carina shorter than posterior ocellus diameter.

NEOTROPICAL TRICHACIS Zootaxa 3337 © 2012 Magnolia Press · 23 FIGURE 10. Bright field images of Trichacis species. A–E: Trichacis hansoni Arias-Penna & Masner, n. sp. A, ♀: habitus, lateral view (holotype); B, head, dorsal view (holotype); C, head, ventrolateral view (holotype); D, ♂, head, anterior view (allotype); E, ♂, metasoma, dorsal view (allotype). F: Trichacis kaulbarsi Arias-Penna & Masner, n. sp., ♂, head, anterior view (allotype).

Mesosoma slightly arched; antero-admedian line present; notaulus absent; anterior region of mesoscutum coriaceous; mesoscutum posteromedially acute; parapsidal line present; central pronotal area smooth; specialized area of mesoscutellum circular and elevated; mesopleural carina incomplete (length of mesopleural carina/length of mesopleuron = 0.29); fore wing distinctly surpassing apex of metasoma. Metasoma moderately elongate; combined length of T3–T6 < T2; T1 shorter than wide; T2 longer than wide; T3–T6 wider than long; anteromedial area of T2 smooth and flattened; T3–T6 densely covered by deep punctuation; T6 subtriangular.

24 · Zootaxa 3337 © 2012 Magnolia Press ARIAS-PENNA ET AL. FIGURE 11. Bright field images of Trichacis species. A–G: Trichacis kaulbarsi Arias-Penna & Masner, n. sp. A, ♂, habitus, dorsal view (allotype); B, ♂, habitus, lateral view (allotype); C, ♂, head and anterior mesosoma, dorsal view (allotype); D, ♂, posterior mesosoma, and metasoma, dorsal view (allotype); E, ♀, head and anterior mesosoma, anterolateral view (holotype); F, ♀, head and anterior mesosoma, dorsal view (holotype); G, ♀, metasoma, dorsal view (holotype). H: Trichacis magnifica Arias-Penna & Masner, n. sp., ♀, head and ventral mesosoma, anterior view (holotype).

NEOTROPICAL TRICHACIS Zootaxa 3337 © 2012 Magnolia Press · 25 Male. Body length: 1.4 mm (allotype);1.60 mm (paratype); mean ± SD = 1.50 ± 0.14 mm. Similar to female except the following: antenna filiform; A1 yellow; A2–A10 brown; leg including coxa yellow; A8 as long as wide; T1 shorter than wide; T2 longer than wide; T3–T5 wider than long; T3–T7 densely covered by deep punctuation. Etymology. Named after M. Kaulbars, the collector of the holotype and allotype. Diagnosis. Trichacis kaulbarsi and T. colombiana have notaulus absent. However, T. kaulbarsi has a smooth interocellar area and a complete hyperoccipital carina. Material examined. Holotype: 1 ♀, MEXICO, SLP 400 km W, Xilitla, 1700 m, June 12 1983, M. Kaulbars (CNCI); allotype: 1 ♂, with the same data as holotype (CNCI); paratype: 1 ♂, with same data as holotype but caught in June 12–August 6 1983, FIT, S. Peck (ICN). Holotype is deposited in CNCI.

Trichacis laticornis Buhl, 2001

Trichacis laticornis Buhl, 2001: 34. Original description.

Diagnosis. Diagnostic characters provided for T. laticornis are temple projection absent; notaulus fading out in anterior fourth; sharp hyperoccipital carina; OOL > LOL; distance between posterior ocellus and hyperoccipital carina shorter than diameter of posterior ocellus; occiput medially smooth and laterovertically striate-coriaceous; mid lobe of mesoscutum posteriorly distinctly blunt, very slightly longer than lateral lobe, nearly (but not quite) reaching base of mesoscutellum; mesopleural carina only visible in slightly less than anterior half of mesopleuron; Tl hardly as long as wide (Buhl 2001, and pers. comm.). The female holotype and the female paratype were collected in Honduras. They were deposited at the Museum of Zoology, Lund University, Sweden (Buhl 2001). Trichacis laticornis is close to T. hansoni but could be distinguished from the latter by its vertex laterovertically striate-coriaceous (coriaceous without striae in T. hansoni).

Trichacis magnifica Arias-Penna & Masner, new species Figures 11H; 12A–12E

Female. Body length: 2.60 mm (holotype); 2.80–3 mm (paratypes, n=2); mean length ± SD = 2.80 ± 0.20 mm. Black; A1 yellow; A2–A3 brown; A5–A10 dark brown; mandible brown with base and apex dark brown; fore and mid legs yellow; hind leg brown; fore and mid coxae brown; hind coxa dark brown; fore wing infuscate. Head, in dorsal view, 2.2 times as wide as long; EH < IOS; interocellar area finely coriaceous; OOL = LOL; antennal clava 5-segmented; A1 surpassing vertex dorsal margin; A2–A10 longer than wide; base of mandible smooth; frons above interantennal process and torulus with transverse striae, with apex Λ-like and reaching, but not surpassing, ventral border of torulus; clypeus exposed; clypeal margin truncate; gena striate; vertex laterally coriaceous and medially smooth; temple projection absent; hyperoccipital carina incomplete: in dorsal view, hyperoccipital carina fading before reaching level of imaginary line connecting eye inner margin to posterior cephalic margin; distance between posterior ocellus and hyperoccipital carina longer than posterior ocellus diameter. Mesosoma slightly arched; antero-admedian line present; notaulus anteriorly incomplete; lateral notaular area with coriaceous sculpture along its anterior margin and inter-notaular area with 1/3 anterior coriaceous; posteromedial region of inter-notaular area convex; parapsidal line present; central pronotal area smooth; specialized area of mesoscutellum cone-like and markedly elevated; mesopleural carina incomplete (length of mesopleural carina/length of mesopleuron = 0.35); fore wing slightly surpassing apex of metasoma. Metasoma moderately elongate; combined length of T3–T6 < T2; T1 slightly longer than wide; T2 longer than wide; T3–T6 wider than long; anteromedial area of T2 smooth and concave; T3–T6 deeply and densely punctured; T6 subtriangular. Male. Body length: 2.40 mm (allotype).

26 · Zootaxa 3337 © 2012 Magnolia Press ARIAS-PENNA ET AL. FIGURE 12. Bright field images of Trichacis magnifica Arias-Penna & Masner, n. sp. A, ♀, habitus, dorsal view (holotype); B, ♀, habitus, lateral view (holotype); C, ♀, head and anterior mesosoma, anterodorsal view (holotype); D, ♂, head and dorsal mesosoma, anterior view (allotype); E, ♂, metasoma, dorsal view (allotype).

NEOTROPICAL TRICHACIS Zootaxa 3337 © 2012 Magnolia Press · 27 Similar to female except the following: A1 yellow; A2–A10 brown; T3 posterior margin smooth and rest densely covered by shallow punctuation; T3–T7 densely covered by deep punctuation. Etymology. The name of the species refers to the splendid specialized area of the mesoscutellum. Diagnosis. Trichacis magnifica is the single species possessing the following combination of characters: OOL = LOL; specialized area of the mesoscutellum cone-like and markedly elevated; hyperoccipital carina incomplete and running until the external margin of posterior ocellus; distance between posterior ocellus and hyperoccipital carina longer than posterior ocellus diameter. Material examined. Holotype: 1 ♀, MEXICO, 20 km W Xititla, 1600 m, June 12 1983, M. Kaulbars (CNCI); allotype: 1 ♂, Aguascalientes, Calvillo, 0.25 mi. S. hwy.70, July 26–August 8 2002, MT, M. Provance (CNCI); paratypes: 1 ♀, Michoacan, 10 mi. S. Uruapan, July 6 1985, SS, J.B. Wooley (CNCI); 1 ♀, with same data as holotype (ICN). Holotype is deposited in CNCI.

Trichacis meridionalis (Brues) 1910

Xestonotus meridionalis Brues 1910. Broteria 9: 150, 151. Original description. Trichacis meridionalis (Brues) 1910. Masner 1965. Comb. n. in: Psyche 1: 303.

Diagnosis. Trichacis meridionalis (Brues 1910) has: antennal scape, pedicel and leg including coxa except tips of hind femur and tibia reddish or ferruginous; vertex and occiput smooth and shining, separated by “a distinct raised line” [= hyperoccipital carina]; ocelli in triangle, lateral ones slightly nearer to eye margin than to median ocellus [= OOL < LOL]; T1 as wide as long. The type was collected in Brazil. Lectotype and paralectotype females were deposited at the Museum of Comparative Zoology, Harvard University, USA. Of the 27 Neotropical species (including those described in this paper), only Trichacis proximata n. sp. and T. acarinata n. sp. have also a smooth vertex. T. acarinata differs from T. meridionalis and T. proximata by hyperoccipital carina absent and T1 longer than wide. T. proximata differs from T. meridionalis by OOL > LOL and T1 shorter than wide.

Trichacis mexicana Arias-Penna & Masner, new species Figures 13A–13F

Female. Body length: 1.70 mm (holotype); 1.60 mm (paratypes, n=2); mean length ± SD = 1.63 ± 0.06 mm. Black; A1–A5 yellow; A6–A10 brown; mandible brown with base black; leg yellow; fore wing infuscate. Head, in dorsal view, 2.5 times as wide as long; EH = IOS; interocellar area with faint, transverse, slightly curved striae; OOL > LOL; antennal clava 4-segmented; A1 not reaching vertex dorsal margin; A2–A4, A10 longer than wide; A5–A6, A9 as long as wide; A7–A8 wider than long; base of mandible smooth; frons above interantennal process and torulus with four well marked transverse striae; interantennal process striate and reaching, but not surpassing, ventral border of torulus; clypeus exposed; clypeal margin truncate; gena striate; vertex laterally coriaceous and medially smooth; temple projection absent; hyperoccipital carina incomplete: in dorsal view, hyperoccipital carina surpassing level of imaginary line connecting eye inner margin to posterior cephalic margin; distance between posterior ocellus and hyperoccipital carina shorter than posterior ocellus diameter. Mesosoma slightly arched; antero-admedian line present but weakly marked; notaulus anteriorly incomplete; lateral notaular area with coriaceous sculpture along its anterior margin; inter-notaular area in its basal 1/3 coriaceous and smooth in its posterior 2/3; posteromedial region of inter-notaular area convex; parapsidal line absent; central pronotal area smooth; specialized area of mesoscutellum heart-shaped; median projection of specialized area of mesoscutellum subtriangular; mesopleural carina incomplete (length of mesopleural carina/ length of mesopleuron = 0.23); fore wing distinctly surpassing apex of metasoma. Metasoma moderately elongate; combined length of T3–T6 < T2; T1 shorter than wide; T2 longer than wide; T3–T5 wider than long; anteromedial area of T2 smooth and flattened; T3–T6 densely covered by deep punctuation; T6 subtriangular. Male. Body length: 2 mm (allotype).

28 · Zootaxa 3337 © 2012 Magnolia Press ARIAS-PENNA ET AL. FIGURE 13. Bright field images of Trichacis mexicana Arias-Penna & Masner, n. sp. A, ♀, head, anterior view (holotype); B, ♀, habitus, dorsal view (holotype); C, ♀, head and anterior mesosoma, dorsal view (holotype); D, ♀, habitus, lateral view (holotype); E, ♂, head and ventral mesosoma, anterolateral view (allotype); F, ♂, posterior mesosoma, and metasoma, dorsal view (allotype).

NEOTROPICAL TRICHACIS Zootaxa 3337 © 2012 Magnolia Press · 29 Similar to female except the following: antenna filiform; A1–A10 yellow; fore and mid legs including coxae yellow; hind leg yellow except apices of femur and tibia brown; hind coxa brown; A5–A9 longer than wide; T1 shorter than wide, T3–T5 smooth along its posterior margin and T6–T7 densely covered by deep punctuation. Etymology. Named after the country of the type locality. Diagnosis. Trichacis mexicana and T. panamana shared the following characters: T1 shorter than wide; clypeus exposed, and clypeal margin truncate. However, T. mexicana has EH = IOS; OOL > LOL and distance between posterior ocellus and hyperoccipital carina shorter than posterior ocellus diameter. Color of antennal segment may vary, for instance holotype has A6–A10 brown and paratype has A7–A10 brown. Material examined. Holotype: 1 ♀, MEXICO. Tamps, Río Frío nr. Gomez Farias, June 6 1983, UV, S. Peck & M. Kaulbars (CNCI); allotype: 1 ♂ with the same data as holotype (CNCI); paratypes: 1 ♀, Tamaulipas 18 m SE Manuel nr. Microondas sta., November 1 1982, SS, J. T. Huber & A. Gonzalez (ICN); 1 ♀ SLP, 1700 m, 40 km W Xilitla, June 12–August 6 1983, S. Peck FIT (CNCI). Holotype is deposited in CNCI.

Trichacis panamana Arias-Penna & Masner, new species Figures 14A–14E; 21D

Female. Body length: 1.80 mm (holotype). Black; A1–A2 yellow; A3–A10 brown; mandible yellow with apex brown; fore and mid legs yellow; hind leg yellow, except tibial apex and tarsi brown; hind coxa brown; fore wing infuscate. Head, in dorsal view, 2.1 times as wide as long; EH < IOS; interocellar area finely coriaceous making weakly transverse striae; OOL < LOL; antenna clava 3-segmented; A1 not reaching vertex dorsal margin; A2–A4, A10 longer than wide; A5–A6, A8 as long as wide; A7–A9 wider than long; base of mandible smooth; frons above torulus with four transverse striae and coriaceous sculpture; interantennal process with reticulate striae, surpassing ventral border of torulus; clypeus exposed; clypeal margin truncate; gena smooth; anterior region of vertex with faint, transverse, slightly curved striae; lateral region of vertex between posterior ocellus and hyperoccipital carina coriaceous, with 3–4 transverse striae reaching external margin of posterior ocellus; lateral region of vertex between hyperoccipital carina and occipital carina coriaceous but without striae; temple projection absent; hyperoccipital carina almost complete: in dorsal view, hyperoccipital carina surpassing level of imaginary line connecting eye inner margin to posterior cephalic margin; distance between posterior ocellus and hyperoccipital carina longer than posterior ocellus diameter. Mesosoma slightly arched; antero-admedian line present; notaulus anteriorly incomplete; lateral notaular area with coriaceous sculpture along its anterior margin; inter-notaular area coriaceous in its basal 1/3 and smooth in its posterior 2/3; posteromedial region of inter-notaular area convex; parapsidal line present; central pronotal area smooth; specialized area of mesoscutellum heart-shaped; median projection of specialized area of mesoscutellum subtriangular; mesopleural carina incomplete (length of mesopleural carina/length of mesopleuron = 0.26); fore wing slightly surpassing apex of metasoma. Metasoma elongate; combined length of T3–T6 < T2; T1 shorter than wide; T2 longer than wide; T3–T6 wider than long; anteromedial area of T2 smooth and flattened; T3 densely covered by shallow punctuation; T4–T6 densely covered by deep punctuation; T6 subtriangular. Male. Body length: 1.90 mm (allotype); 1.80 mm (paratype); mean length ± SD = 1.85 ± 0.07 mm. Similar to female except the following: hind leg yellow except tibial apex and tarsi dark brown; antenna filiform; A1–A3 yellow; A4–A10 brown; A3 as long as wide; A5–A9 longer than wide; T1 shorter than wide; T3 smooth along its posterior margin and T4–T6 covered by deeply and dense punctuation. Etymology. Named after the country of the type locality. Diagnosis. Both Trichacis panamana and T. mexicana have: T1 shorter than wide; clypeus exposed, and clypeal margin truncate. However, T. panamana has EH < IOS, OOL < LOL and distance between posterior ocellus and hyperoccipital carina longer than posterior ocellus diameter. Material examined. Holotype: 1 ♀, PANAMA, Veraguas, 8 km W Santa Fe, Forest, 8º30´26´´N 81º06´49´´W, 870 m, January 1–4 2001, YPT, L. Masner (CNCI); allotype: 1 ♂, with the same data as holotype (CNCI); paratype: 1 ♂, with the same data as holotype (ICN). Holotype is deposited in CNCI.

30 · Zootaxa 3337 © 2012 Magnolia Press ARIAS-PENNA ET AL. FIGURE 14. Bright field images of Trichacis species. A–E: Trichacis panamana Arias-Penna & Masner, n. sp., A, ♀, head and ventral mesosoma, anterior view (holotype); B, ♀, habitus, dorsal view (holotype); C, ♀, habitus, lateral view. D, ♀, head, dorsal view (holotype). E, ♂, head and ventral mesosoma, anterior view (allotype). F–H, Trichacis pecki Arias-Penna & Masner, n. sp., ♀ (holotype). F, head, anterior view; G, habitus, dorsal view; H, habitus, lateral view.

NEOTROPICAL TRICHACIS Zootaxa 3337 © 2012 Magnolia Press · 31 Trichacis pecki Arias-Penna & Masner, new species Figures 14F–14H; 15A, 15B

Female. Body length: 1.30 mm (holotype);1.50 mm (paratypes, n=3); mean length ± SD = 1.45 ± 0.10 mm. Black; A1–A7 yellow; A8–A10 brown; mandible brown with apex black; fore and mid legs yellow; hind leg with trochanter and basal 1/3 of femur yellow, otherwise dark brown; hind coxa yellow; fore wing infuscate. Head, in dorsal view, 2.4 times as wide as long; EH < IOS; interocellar area finely coriaceous; OOL > LOL; antennal clava 3-segmented; A1 not reaching vertex dorsal margin; A2, A5–A7, A10 longer than wide; A3–A4 as long as wide; A8–A9 wider than long; base of mandible smooth; frons above torulus with three transverse striae; interantennal process smooth, curving down and surpassing ventral border of torulus; clypeus exposed; clypeal margin convex medially; gena smooth; lateral region of vertex between posterior ocellus and hyperoccipital carina weakly coriaceous, with cluster of 4-6 well-marked longitudinal curved striae, which seem to emerge from hyperoccipital carina; lateral region of vertex between hyperoccipital carina and occipital carina coriaceous; temple projection absent; hyperoccipital carina incomplete but curving up and merging with stria running along the eye inner margin; distance between posterior ocellus and hyperoccipital carina longer than posterior ocellus diameter. Mesosoma arched; antero-admedian line present; notaulus anteriorly incomplete; lateral notaular area with coriaceous sculpture along its anterior margin; inter-notaular area coriaceous in its basal 1/3 and smooth in its posterior 2/3; posteromedial region of inter-notaular area acute; parapsidal line present; central pronotal area with anterior region smooth and rest coriaceous; specialized area of mesoscutellum heart-shaped; median projection of specialized area of mesoscutellum subtriangular; mesopleural carina incomplete (length of mesopleural carina/ length of mesopleuron = 0.26); fore wing slightly surpassing apex of metasoma. Metasoma moderately elongate; combined length of T3–T6 < T2; T1 shorter than wide; T2 longer than wide; T3–T6 wider than long; anteromedial area of T2 smooth and flattened; T3 densely covered by shallow punctuation, T4–T6 densely covered by deep punctuation; T6 subtriangular. Male. Unknown. Etymology. Named after S. & J. Peck, the collectors of numerous specimens of Trichacis. Diagnosis. Trichacis pecki, T. fernandezi and T delsinnei have an incomplete hyperoccipital carina, curving up when reaching eye margin. The sculpture of vertex and the acute shape of the posteromedial region of inter- notaular area allow to distinguish T. pecki from these species. Material examined. Holotype: 1 ♀, ECUADOR, Esm., Bilsa Biol. Sta., 0º34´N 79º71´W, 500 m, May 10–June 4 1996, MT, P. Hibbs (CNCI); paratypes: 3 ♀♀, Pinch. 16 km E, Santo Domingo, Tinalandia, 680 m, May 4–July 25 1985, MT–FIT, S. & J. Peck (2 at CNCI and 1 at ICN). Holotype is deposited in CNCI.

Trichacis procera Arias-Penna & Masner, new species Figures 15C–15F

Female. Body length: 2.40 mm (holotype). Black shining; A1–A5 yellow, A6–A10 brown; mandible brown with apex dark brown; leg yellow; fore wing infuscate. Head, in dorsal view, 2.4 times as wide as long; EH < IOS; interocellar area smooth; OOL > LOL; antennal clava 4-segmented; A1 not reaching vertex dorsal margin; A2–A4, A7, A10 longer than wide; A5, A6, A8, A9 wider than long; base of mandible smooth; frons above interantennal process and torulus with three well-marked transverse striae; interantennal process smooth, curving down and surpassing ventral border of torulus; clypeus hidden by interantennal process; clypeal margin truncate; gena smooth; lateral region of vertex between posterior ocellus and hyperoccipital carina weakly coriaceous; temple projection absent; hyperoccipital carina complete; distance between posterior ocellus and hyperoccipital carina shorter than posterior ocellus diameter. Mesosoma slightly arched; antero-admedian line present; notaulus anteriorly incomplete; mesoscutum smooth; posteromedial region of inter-notaular area convex; parapsidal line present; central pronotal area smooth; specialized area of mesoscutellum heart-shaped; median projection of specialized area of mesoscutellum triangular and with a broad tuft of hairs; mesopleural carina complete; fore wing slightly surpassing apex of metasoma.

32 · Zootaxa 3337 © 2012 Magnolia Press ARIAS-PENNA ET AL. FIGURE 15. Bright field images of Trichacis species. A, B: Trichacis pecki Arias-Penna & Masner, n. sp., ♀ (holotype). A, head and anterior mesosoma, dorsal view; B, head and mesosoma, dorsolateral view; C–F: Trichacis procera Arias-Penna & Masner, n. sp., ♀ (holotype). C, head, anterior view; D, head and mesosoma, dorsal view; E, metasoma, dorsolateral view; F, head and mesosoma, lateral view.

NEOTROPICAL TRICHACIS Zootaxa 3337 © 2012 Magnolia Press · 33 Metasoma elongate; combined length of T3–T6 < T2; T1, T3–T4, T6 longer than wide; T2, T5 shorter than wide; anteromedial area of T2 smooth and concave; T3 smooth along its anterior margin and densely covered by shallow punctuation; T4–T6 densely covered by deep punctuation; T6 subtriangular. Male. Unknown. Etymology. The name of the species refers to its long interantennal process. Diagnosis. Trichacis procera and T. dianae have a complete mesopleural carina but T. procera has OOL > LOL; lateral region of its vertex is weakly coriaceous between posterior ocellus and hyperoccipital carina. Material examined. Holotype: 1 ♀, MEXICO, Monte Bello, Chiapas, July 15–31 1969, MT, W. Mason (CNCI). Holotype is deposited in CNCI.

Trichacis proximata Arias-Penna & Masner, new species Figures 16A–16E; 21B

Female. Body length: 2.30 mm (holotype); 2.05 mm (paratype); mean length ± SD = 2.18 ± 0.18 mm. Black and shining; A1–A6 yellow; A7–A10 dark brown; mandible black with apex brown; fore leg yellow; mid leg brown, except dark brown patches on femur medially, and on tibial apex; hind leg with apical 2/3 of femur, tibia and tarsi bicolored, dark brown and black, otherwise brown; fore coxa brown; hind coxa dark brown; fore wing infuscate except in basal region and margin. Head, in dorsal view, 2.7 times as wide as long; EH < IOS; interocellar area smooth; OOL > LOL; antennal clava 4-segmented; A1 not reaching vertex dorsal margin; A2–A5, A10 longer than wide; A6 wider than long; A7–A9 as long as wide; base of mandible striate; frons mostly smooth, with three transverse striae above torulus, and with scattered short pilosity in its anterior region (at high magnification [e.g. 160x] this gives impression of punctuation); interantennal process smooth and curving down, surpassing ventral border of torulus; clypeus hidden by interantennal process; clypeal margin convex medially; gena striate; vertex smooth; temple projection absent; hyperoccipital carina complete; posterior ocellus reaching hyperoccipital carina. Mesosoma slightly arched; antero-admedian line absent; notaulus anteriorly incomplete; lateral notaular area smooth and inter-notaular area with basal 1/3 coriaceous and posterior 2/3 smooth; posteromedial region of inter- notaular area convex; parapsidal line present; central pronotal area smooth; specialized area of mesoscutellum heart-shaped; median projection of specialized area of mesoscutellum triangular; mesopleural carina incomplete (length of mesopleural carina/length of mesopleuron = 0.57); fore wing slightly surpassing apex of metasoma. Metasoma elongate; combined length of T3–T6 < T2; T1 shorter than wide; T2 longer than wide; T3–T5 shorter than wide; anteromedial area of T2 smooth and concave; T3 mostly smooth, with shallow punctuation along its posterior border; T4 smooth along its anterior margin and T5–T6 densely covered by deep punctuation; T6 subtriangular, almost as long as wide. Male. Unknown. Etymology. The name of the species refers to the close position of the posterior ocelli relative to the hyperoccipital carina. Diagnosis. Trichacis proximata is the single species that presents the posterior ocelli close to the hyperoccipital carina. Material examined. Holotype: 1 ♀, COSTA RICA, Guan. GCA, Est. Maritza, 700 m, January 20–February 3 1996, MT/YPT, J.S. Noyes (BMNH); paratype: 1 ♀, Guan. GCA, Est. Pitilla, 700 m, February 12–27 1996, MT/ YPT, J.S. Noyes (CNCI). Holotype is deposited in CNCI.

Trichacis punctata Arias-Penna & Masner, new species Figures 16F; 17A–17E

Female. Body length: 2.30 mm (holotype). Black; mandible yellow with apex brown; A1 yellow; A2–A10 brown; fore and mid legs yellow; hind leg with tarsi and apices of femur and tibia brown, otherwise yellow; hind coxa dark brown; fore wing infuscate in posterior 2/3, with darker and lighter patterns in basal 1/3.

34 · Zootaxa 3337 © 2012 Magnolia Press ARIAS-PENNA ET AL. FIGURE 16. Bright field images of Trichacis species. A–E: Trichacis proximata Arias-Penna & Masner, n. sp., ♀ (holotype). A, head, anterior view; B, habitus, dorsal view; C, habitus, lateral view; D, head and anterior mesosoma, dorsal view; E, posterior mesosoma and metasoma, dorsal view. F: Trichacis punctata Arias-Penna & Masner, n. sp., ♀, head and anteroventral mesosoma, anterior view (holotype).

NEOTROPICAL TRICHACIS Zootaxa 3337 © 2012 Magnolia Press · 35 FIGURE 17. Bright field images of Trichacis species. A–E: Trichacis punctata Arias-Penna & Masner, n. sp., ♀ (holotype). A, habitus, dorsal view; B, habitus, lateral view; C, head, posterodorsal view; D, head, dorsal view; E, posterior head, mesosoma and anterior metasoma, dorsal view. F–H: Trichacis sculpturata Arias-Penna & Masner, n. sp., ♀ (holotype). F, head and ventral mesosoma, anterior view; G, habitus, dorsal view; H, habitus lateral view.

36 · Zootaxa 3337 © 2012 Magnolia Press ARIAS-PENNA ET AL. Head, in dorsal view, 2.2 times as wide as long; EH < IOS; interocellar area smooth; OOL > LOL; antennal clava 4-segmented; A1 not reaching vertex dorsal margin; A2–A4, A6, A10 longer than wide; A5, A7–A9 as long as wide; base of mandible smooth; frons above torulus with transverse striae; interantennal process reaching, but not surpassing, ventral border of torulus; clypeus exposed; clypeal margin truncate; gena striate; vertex with scattered punctures laterally; temple projection absent; hyperoccipital carina absent; length between posterior ocellus and occipital carina line > OOL. Mesosoma arched; antero-admedian line present; notaulus anteriorly incomplete; lateral notaular area smooth; inter-notaular area with anterior 1/3 coriaceous; posteromedial region of inter-notaular area convex; parapsidal line present; central pronotal area smooth; specialized area of mesoscutellum heart-shaped; median projection of specialized area of mesoscutellum rectangular; mesopleural carina incomplete (length of mesopleural carina/length of mesopleuron = 0.50); fore wing distinctly surpassing apex of metasoma. Metasoma elongate; combined length of T3–T6 < T2; T1 shorter than wide; anteromedial area of T2 smooth and flattened; T2 longer than wide; T3–T5 densely covered by deep punctuation; T6 densely covered by deep punctuation along its anterior border. Male: Unknown. Etymology. The name refers to the punctures present on vertex. Diagnosis. Trichacis punctata and T. acarinata are characterized by the absence of hyperoccipital carina. T. punctata can be distinguished by the combination of the following characters: combined length of T3–T6 < T2; posteromedial region of inter-notaular area convex; fore wing distinctly surpassing apex of metasoma. Material examined. Holotype: 1 ♀, BRAZIL, S. Bocaina, S. José Barre S.P., 1650 m, November 1968, M. Alvarenga (CNCI). Holotype is deposited in CNCI.

Trichacis sculpturata Arias-Penna & Masner, new species Figures 17F–17H; 18A

Female. Body length: 2.25 mm (holotype). Black; A1–A2 yellow; A3–A10 brown; base of mandible black, apex brown and rest yellow; fore and mid legs yellow; hind leg yellow but with tibial apex and tarsi brown; hind coxa brown; fore wing infuscate. Head, in dorsal view, 1.8 times as wide as long; EH < IOS; interocellar area finely coriaceous; OOL > LOL; antennal clava 4-segmented; A1 not reaching vertex dorsal margin; A2–A7, A10 longer than wide; A8–A9 as long as wide; base of mandible striate; frons above torulus and interantennal process coriaceous with three marked transverse striae; interantennal process smooth and reaching, but not surpassing, ventral border of torulus; clypeus exposed; clypeal margin convex medially; gena striate; vertex laterally coriaceous; medial vertex area smooth between posterior ocellus and hyperoccipital carina and coriaceous between hyperoccipital carina and occipital carina; temple projection absent; hyperoccipital carina incomplete: in dorsal view, hyperoccipital carina fading before reaching level of imaginary line connecting eye inner margin to posterior cephalic margin; distance between posterior ocellus and hyperoccipital carina longer than posterior ocellus diameter. Mesosoma arched; antero-admedian line present; notaulus percurrent; lateral notaular area with coriaceous sculpture along its anterior margin; inter-notaular area coriaceous; posteromedial region of inter-notaular area convex; parapsidal line present; central pronotal area with anterior margin and rest coriaceous; specialized area of mesoscutellum heart-shaped; median projection of specialized area of mesoscutellum triangular; mesopleural carina incomplete (length of mesopleural carina/length of mesopleuron = 0.48); fore wing distinctly surpassing apex of metasoma. Metasoma elongate; combined length of T3–T6 < T2; T1 shorter than wide; T2 longer than wide; anteromedial area of T2 concave and covered with irregular striae, T3 densely covered by shallow punctuation, T4–T6 densely covered by deep punctuation; T6 subtriangular. Male. Unknown. Etymology. The name of the species refers to the coriaceous sculpture of the inter-notaular area. Diagnosis. Trichacis sculpturata and T. corrugata have inter-notaular area coriaceous. T. sculpturata differs by the sculpture of its vertex and by the shape of its specialized area of the mesoscutellum. Material examined. Holotype: 1 ♀, MEXICO, Chis. 7200 ft, S. Crist., Las Casas, July 1–12 1969, MT (CNCI). Holotype is deposited in CNCI.

NEOTROPICAL TRICHACIS Zootaxa 3337 © 2012 Magnolia Press · 37 FIGURE 18. Bright field images of Trichacis species. A: Trichacis sculpturata Arias-Penna & Masner, n. sp., ♀, head, dorsal view (holotype). B–E: Trichacis transversata Arias-Penna & Masner, n. sp., ♀ (holotype). B, head, anterior view; C, habitus, dorsal view; D, habitus, lateral view; E, head and mesosoma, dorsolateral view. F–H: Trichacis triangulata Arias-Penna & Masner, n. sp., ♂ (allotype). F, head and ventral mesosoma, anterior view; G, habitus, dorsal view; H, head and mesosoma, dorsal view.

38 · Zootaxa 3337 © 2012 Magnolia Press ARIAS-PENNA ET AL. Trichacis transversata Arias-Penna & Masner, new species Figures 18B–18E

Female. Body length: 2.20 mm (holotype); 2–2.20 mm (paratypes, n=2); mean length ± SD = 2.13 ± 0.12 mm. Black; radicle yellow, base of A1 yellow, distal part of A1 and A2–A10 brown; mandible yellow with apex brown; fore leg yellow; mid leg yellow but with femur apex and tarsi dark brown; hind leg with trochanter yellow, bases of femur and tibia brown and rest dark brown; hind coxa dark brown; fore wing infuscate. Head, in dorsal view, 2.07 times as wide as long; EH < IOS; interocellar area finely coriaceous and with shallow transverse striae; OOL > LOL; antennal clava 3-segmented; A1 not reaching vertex dorsal margin; A2–A6, A10 longer than wide; A7–A9 as long as wide; base of mandible smooth; frons above torulus weakly coriaceous and with seven transverse striae; clypeus exposed; clypeal margin convex medially; interantennal process surpassing ventral border of torulus; gena striate; lateral region of vertex between posterior ocellus and hyperoccipital carina coriaceous, and between hyperoccipital carina and occipital carina with well-marked longitudinal striae; temple projection absent; hyperoccipital carina almost complete: in dorsal view, hyperoccipital carina surpassing level of imaginary line connecting eye inner margin to posterior cephalic margin; distance between posterior ocellus and hyperoccipital carina shorter than posterior ocellus diameter. Mesosoma arched; antero-admedian line absent; notaulus anteriorly incomplete; lateral notaular area smooth; inter-notaular area with basal 1/3 coriaceous and posterior 2/3 smooth; posteromedial region of inter-notaular area broadly convex; parapsidal line present; central pronotal area smooth; specialized area of mesoscutellum heart- shaped; median projection of specialized area of mesoscutellum subtriangular; mesopleural carina incomplete (length of mesopleural carina/length of mesopleuron = 0.13); fore wing distinctly surpassing apex of metasoma. Metasoma elongate; combined length of T3–T6 < T2; T1–T2 longer than wide; T3–T5 wider than long; T6 shorter than wide; anteromedial area of T2 smooth and flattened; T3 densely covered by shallow punctuation; T4–T5 and T6 along its anterior border, densely covered by deep punctuation; T6 subtriangular. Male. Unknown. Etymology. The name of the species refers to the fine transverse striae located between the anterior and posterior ocelli. Diagnosis. Trichacis transversata is the single species with the combination of the following characters: interocellar area finely coriaceous forming fine transverse striae; lateral region of vertex between posterior ocellus and hyperoccipital carina coriaceous, and between hyperoccipital carina and occipital carina with well-marked longitudinal striae. Material examined. Holotype: 1 ♀, COSTA RICA, Guan., Cerro de la Muerte, 4 km NE Canon, 2350 m, January 1995, MT, P. Hanson (CNCI); paratypes: 1 ♀, Cartago, 4 km NE Canon, MT, Genesis, 2350 m, February- July 1995, P. Hanson (CNCI); 1 ♀, with same data as holotype (ICN). Holotype is deposited in CNCI.

Trichacis triangulata Arias-Penna & Masner, new species Figures 18F–18H; 19; 20A–20F

Female. Body length: 2.05 mm (holotype); 1.95–2.20 mm (paratypes, n=3); mean length ± SD = 2.09 ± 0.11 mm. Black; A1–A10 brown; mandible black with apex brown; fore leg yellow; mid leg brown, with apical ½ of tibia darker; hind leg brown, with tarsi and apical ½ of femur and tibia darker; fore coxa brown; hind coxa dark brown, fore wing infuscate. Head, in dorsal view, 2.6 times as wide as long; EH < IOS; interocellar area finely coriaceous; OOL > LOL; antennal clava 3-segmented; A1 not reaching vertex dorsal margin; A2–A4 and A10 longer than wide; A5–A6 as long as wide; A7–A9 wider than long; base of mandible smooth; frons above torulus and interantennal process coriaceous with three marked transverse striae; interantennal process surpassing ventral margin of torulus; clypeus exposed; clypeal margin convex medially; gena striate; vertex with lateral region coriaceous and with medial region smooth between posterior ocellus and hyperoccipital carina and coriaceous between hyperoccipital carina and occipital carina; temple projection absent; hyperoccipital carina complete; distance between posterior ocellus and hyperoccipital carina shorter than posterior ocellus diameter.

NEOTROPICAL TRICHACIS Zootaxa 3337 © 2012 Magnolia Press · 39 FIGURE 19. Trichacis triangulata Arias-Penna & Masner, n. sp., ♂, habitus, lateral view (allotype).

Mesosoma arched; antero-admedian line present; notaulus anteriorly incomplete; lateral notaular area with coriaceous sculpture along its anterior margin; inter-notaular area with basal 1/3 coriaceous and posterior 2/3 smooth; posteromedial region of inter-notaular area convex; parapsidal line present; central pronotal area with posterior half coriaceous; specialized area of mesoscutellum with broad triangular sheet-like projection, moderately elevated; mesopleural carina incomplete (length of mesopleural carina/length of mesopleuron = 0.29); fore wing distinctly surpassing apex of metasoma.

40 · Zootaxa 3337 © 2012 Magnolia Press ARIAS-PENNA ET AL. FIGURE 20. Trichacis triangulata Arias-Penna & Masner, n. sp. A–C: ♂ (allotype). A, head and anterior mesosoma, lateral view; B, metasoma, dorsal view; C, head, dorsal view. D–F: ♀ (holotype). D, head and ventral mesosoma, anterior view; E, head and mesosoma, dorsolateral view; F, metasoma, dorsal view. Metasoma elongate; combined length of T3–T6 < T2; T1 shorter than wide; T2 slightly longer than wide; anteromedial area of T2 smooth and flattened; T3 anterior margin smooth and with shallow punctuation; T4–T6 densely covered by deep punctuation; T6 subtriangular. Male. Body length: 2.20 mm (allotype); 1.95–2.30 mm (paratypes, n=10); mean length ± SD = 2.09 ± 0.11 mm. Similar to female except the following: head, in dorsal view, twice as wide as long; EH < IOS; antenna filiform; A5, A7–A9 longer than wide; T1–T2 longer than wide.

NEOTROPICAL TRICHACIS Zootaxa 3337 © 2012 Magnolia Press · 41 Etymology. The name of the species refers to the shape of its specialized area of the mesoscutellum. Diagnosis. Trichacis triangulata is the only species to have the combination of the following characters: distance between posterior ocellus and hyperoccipital carina shorter than posterior ocellus diameter; T1 shorter than wide, and specialized area of mesoscutellum with broad triangular sheet-like projection. Material examined. Holotype: 1 ♀, MEXICO, Oax, 26 km E Valle Nacional, 1220 m, June 25 1983, M. Kaulbars & S. Peck; allotype:1 ♂, with same data as holotype (CNCI); paratypes: 9 ♂♂, 3 ♀♀, with same data as holotype (CNCI), 1 ♂, with same data as holotype but collected by S. Peck (ICN). Holotype is deposited in CNCI.

FIGURE 21. Variation observed in extent and shape of the hyperoccipital carina. A–E. A, hyperoccipital carina absent (Trichacis acarinata Arias-Penna & Masner, n. sp. ♀ holotype). B, complete, reaching the eye margin (Trichacis proximata Arias-Penna & Masner, n. sp. ♀ holotype). C, incomplete, fading before reaching level of imaginary line (= the white line on the figure) connecting eye inner margin to posterior cephalic margin (Trichacis acuta Arias-Penna & Masner, n. sp. ♀ holotype). D, incomplete but surpassing level of imaginary line (Trichacis panamana Arias-Penna & Masner, n. sp. ♀ holotype). E, incomplete, surpassing level of imaginary line, curving up and merging with stria running along the eye inner margin (Trichacis fernandezi Arias-Penna & Masner, n. sp. ♀ holotype). White arrows indicate the extremities of the hyperoccipital carina when present.

42 · Zootaxa 3337 © 2012 Magnolia Press ARIAS-PENNA ET AL. Acknowledgements

We are very grateful to J. Huber and A. Bennett, curators at the CNCI, for the loan of specimens. P. N. Buhl kindly shared useful information and answered swiftly to all our questions about the Trichacis species he described. M. Leponce (RBINS) allowed us to use the camera of his laboratory. Y. Laurent provided some helpful advice on digitalization. I. Bachy improved the image quality with Adobe Photoshop CS. We thank two anonymous reviewers and I. Mikó for their valuable comments. We thank M. Sharkey (University of Kentucky), F. Fernández (ICN) and D. C. Arias-Penna (University of Illinois) for their unconditional help. This study was made possible by financial support in part from the National Science Foundation under grant No. DEB 0205982 and from CanaCol Foundation (2004) grant to TMAP.

References

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44 · Zootaxa 3337 © 2012 Magnolia Press ARIAS-PENNA ET AL. APPENDIX 1. Measurements in millimeter of primary types (holotype: ♀, allotype: ♂) of the nominal species of Trichacis from South and Central America. Abbreviations: BL=body length, HW=maximum head width (dorsal view), HL=maximum head length (dorsal view, head dorsoventral axis is perpendicular to longitudinal body axis), EH=maximum eye height (anterior view), IOS=interorbital space (shortest distance between eyes, anterior view), POL=posterior ocellar line (shortest distance between posterior ocelli), OOL=ocular ocellar line (the shortest distance between the posterior ocellus and the eye, LOL=lateral ocellar line (the shortest distance between the posterior and anterior ocelli), A1–A10L=antennomere 1–10 length (shortest distance between the proximal and distal margins of the antennomere), antennomeres 1–10 width (shortest distance between the lateral margins of the antennomere), POOCL=posterior ocellus–occipital carina line (shortest distance between the posterior ocellus and the occipital carina, not applicable if the hyperoccipital carina is present), POHOCL=posterior ocellus–hyperoccipital carina line (shortest distance between the posterior ocellus and hyperoccipital carina, not applicable if the hyperoccipital carina is absent), OHOCL=ocular–hyperoccipital carina length (shortest distance between the eye and the hyperoccipital carina, not applicable if the hyperoccipital carina is absent), AALL=antero-admedian line length (shortest distance between the anterior and posterior margin of the antero-admedian line in dorsal view), PLL=parapsidal line length (shortest distance between the anterior and posterior margins of the parapsidal line in dorsal view), CPAL=central pronotal area length (shortest distance between the anterior and posterior margins of the pronotum in dorsal view), CPAW=central pronotal area width (shortest distance between the dorsal most points of the epomiae in dorsal view), MCL=mesopleural carina length (shortest distance between the anterior and posterior margin of the mesopleural carina in lateral view), PL2l=mesopleuron length (shortest distance between the anterior and posterior margins of the mesopleuron in lateral view), MTSL=metasoma length (shortest distance between the anterior and posterior margins of the metasoma in dorsal view), MTSW=metasoma width (longest distance between the lateral margins of the metasoma that is perpendicular to the anteroposterior body axis in dorsal view), MTSH=metasoma height (the longest distance between the dorsal and ventral margins of the metasoma that is vertical and perpendicular to the anteroposterior body axis), T1–T7L=metasoma terga 1–7 length (the length of the longitudinal median line of the metasomal tergum), T1–T7W=metasomal terga 1–7 width (the longest distance between the lateral margins of the metasomal tergum in dorsal view).

.

NEOTROPICAL TRICHACIS Zootaxa 3337 © 2012 Magnolia Press · 45 1.9 (2) (2) 1.9 (0.52) 0.54 (0.27) 0.2 (0.26) 0.28 (0.27) 0.29 (0.96) 0.97 (0.13) 0.1 (0.056) 0.06 (0.068) 0.07 (0.82) 0.9 2.6 (2.4) (2.4) 2.6 (0.65) 0.66 (0.3) 0.3 (0.31) 0.31 (0.36) 0.4 (0.86) 0.77 (0.17) 0.29 (0.075) 0.09 (0.075) 0.09 (1) 1 1.95 0.52 0.30 0.23 0.38 0.6 0.18 0.1 0.1 0.18 0.6 0.38 0.08 1.25 0.23 0.30 0.52 1.95 0.88 0.09 0.08 0.19 1.71 0.07 0.12 0.82 0.16 0.39 0.86 0.32 0.37 0.3 0.65 2.6 0.32 0.35 0.67 2.2 0.87 0.08 0.07 0.16 0.87 0.33 0.29 0.27 0.55 1.6 2.25 0.18 0.65 0.86 0.36 0.09 0.31 0.35 1.3 0.069 1.8 (1.9) (1.9) 1.8 (0.53) 0.54 (0.31) 0.25 (0.28) 0.23 (0.29) 0.31 (0.96) 0.74 (0.18) 0.18 (0.075) 0.05 (0.09) 0.08 (0.83) 0.62 2.2 (2.2) (2.2) 2.2 (0.63) 0.65 (0.31) 0.3 (0.31) 0.31 (0.39) 0.39 (0.8) 0.8 (0.19) 0.17 (0.12) 0.11 (0.08) 0.09 (1.5) 1.2 2.3 0.6 0.27 0.29 0.35 0.82 0.17 0.11 0.07 1.6 1.6 0.07 0.11 0.17 0.82 0.35 0.29 0.6 2.3 0.27 0.06 0.13 0.075 1.25 2.4 1.43 0.28 0.4 0.45 0.21 1.3 0.06 0.08 0.13 0.89 0.28 0.25 0.5 1.3 0.21 0.11 0.18 0.9 0.07 1.6 0.33 0.29 2.2 0.27 0.56 2.5 (1.9) (1.9) 2.5 (0.5) 0.55 (0.26) 0.27 (0.23) 0.26 (0.27) 0.33 (0.85) 0.78 (0.18) 0.16 (0.11) 0.07 (0.087) 0.04 (1.3) 1.75 ) ) (2.2) 2.05 0.6 (0.63) (0.23) 0.23 (0.29) 0.27 (0.34) 0.32 (0.85) 0.8 (0.16) 0.17 (0.09) 0.08 (0.075) 0.07 (1.2) 1.14 ) (1.9) 1.8 (0.52) 0.49 (0.25) 0.24 (0.25) 0.25 (0.31) 0.3 (0.8) 0.8 (0.16) 0.15 (0.069) 0.068 (0.062) 0.06 (1.1) 1.1 ) ) ) ) (2) 1.7 (0.6) 0.5 (0.25) 0.2 (0.3) 0.3 (0.3) 0.3 1(1) (0.17) 0.14 (0.09) 0.09 (0.07) 0.07 (1.3) 1.3 ) (1.4) 1.6 (0.49) 0.44 (0.25) 0.23 (0.22) 0.21 (0.32) 0.32 (0.69) 0.65 (0.14) 0.12 (0.94) 0.11 (0.125) 0.06 (1.5) 1.83 ♂ ♂ ♂ ) ) (2) 2.5 (0.59) 0.62 (0.24) 0.25 (0.29) 0.31 (0.31) 0.35 (0.93) 0.88 (0.17) 0.14 (0.11) 0.15 (0.069) 0.068 (1.6) 2.2 ♂ ) ( ♂ ♂ ♀ ( ( 1.4 1.4 0.13 0.07 0.5 0.86 0.28 0.07 0.24 0.23 1 2.3 0.14 0.13 0.7 0.4 0.09 0.085 0.34 0.26 1.4 ) ( ♂ ♀ ♀ 2.8 0.61 0.3 0.28 0.27 0.14 0.93 0.081 1.17 0.069 ( ♀ ( ♂ ♀ ) 2.3 0.61 0.34 0.31 0.33 0.94 0.18 0.08 0.09 0.89 0.89 0.08 0.09 0.18 0.94 0.33 0.31 0.34 2.3 0.61 ♀ ♀ ♀ ( ♀ ♀ ♂ ♀ ♀ ( 1.05 0.2 0.12 0.19 0.77 0.52 0.4 0.35 0.85 2.4 ♀ ♀ ♀ ♀ ♂ ♀ ♀ ( ♀ ♀ ♀ ( ♀ n. sp. n. sp. sp. n. sp. n. ♀ n. sp. sp. n. ♀ n. sp. sp. n. n. sp. sp. n. sp. n. sp. n. n. sp. sp. n. sp. n. n. sp. sp. n. sp. n. n. sp. n. sp. n. sp. n. sp. n. sp. sp. n. n. sp. n. n. sp. sp. n. n. sp. sp. n. n. sp. n. sp. n. sp. n. sp. n. sp. n. Species Species POLOOL BL LOL OOL/LOL HW HL EH IOST. acarinata EH/IOS acuminata T. T. acuta clypeata T. T. colombiana T. concavata T. corrugata T. costaricana delsinnei T. T. depressa T. dianae fernandezi T. T. hansoni T. kaulbarsi magnifica T. T. mexicana panamana T. T. pecki procera T. proximata T. T. punctata sculpturata T. transversata T. T. triangulata

46 · Zootaxa 3337 © 2012 Magnolia Press ARIAS-PENNA ET AL. ) 0.28 (0.3) (0.3) 0.28 (0.06) 0.075 (0.04) 0.075 (0.075) 0.04 (0.05) 0.075 (0.04) 0.05 (0.1) 0.05 (0.04) 0.05 (0.07) 0.06 0.45 (0.4) (0.4) 0.45 (0.05) 0.06 (0.09) 0.1 (0.04) 0.05 (0.06) 0.11 (0.05) 0.04 (0.14) 0.11 (0.05) 0.04 (0.11) 0.1 0.36 0.36 0.06 0.037 0.044 0.044 0.047 0.037 0.087 0.069 0.4 0.04 0.03 0.07 0.04 0.09 0.1 0.04 0.26 0.05 0.06 0.05 0.03 0.04 0.04 0.04 0.06 0.075 0.4 0.05 0.06 0.05 0.08 0.05 0.06 0.11 0.075 0.43 0.05 0.06 0.05 0.08 0.05 0.06 0.08 0.08 0.66 0.05 0.09 0.04 0.08 0.05 0.06 0.09 0.11 0.42 0.1 0.04 0.05 0.08 0.09 0.05 0.09 0.06 0.44 0.04 0.08 0.09 0.04 0.09 0.1 0.05 0.07 0.41 (0.43) (0.43) 0.41 (0.06) 0.06 (0.075) 0.1 (0.05) 0.04 (0.075) 0.1 (0.05) 0.04 (0.09) 0.06 (0.05) 0.04 (0.06) 0.07 0.37 0.37 0.1 0.05 0.04 0.07 0.09 0.1 0.07 0.04 0.37 0.03 0.05 0.07 0.04 0.09 0.1 0.04 0.07 0.32 0.03 0.04 0.04 0.06 0.04 0.04 0.07 0.09 0.26 0.04 0.04 0.04 0.04 0.04 0.04 0.05 0.07 0.45 0.05 0.09 0.06 0.04 0.07 0.1 0.05 0.04 0.36 0.04 0.08 0.04 0.07 0.05 0.05 0.06 0.075 0.33 (0.34) (0.34) 0.33 (0.075) 0.07 (0.08) 0.09 (0.04) 0.04 (0.04) 0.06 (0.04) 0.04 (0.08) 0.05 (0.04) 0.04 (0.06) 0.04 0.5 (0.4) (0.4) 0.5 (0.05) 0.07 (0.1) 0.11 (0.04) 0.04 (0.06) 0.11 (0.04) 0.04 (0.1) 0.1 (0.05) 0.04 (0.04) 0.06 ) (0.4) 0.39 (0.06) 0.07 (0.09) 0.09 (0.04) 0.04 (0.05) 0.08 (0.03) 0.04 (0.11) 0.09 (0.05) 0.05 (0.06) 0.05 ) (0.35) 0.31 (0.07) 0.06 (0.069) 0.07 (0.04) 0.04 (0.05) 0.07 (0.04) 0.04 (0.1) 0.08 (0.04) 0.04 (0.04) 0.06 ) ) ) (0.37) 0.34 (0.06) 0.06 (0.1) 0.05 (0.05) 0.04 (0.06) 0.05 (0.05) 0.04 (0.1) 0.06 (0.05) 0.04 (0.06) 0.04 ) (0.31) 0.32 (0.05) 0.06 (0.07) 0.08 (0.04) 0.04 (0.07) 0.06 (0.05) 0.04 (0.06) 0.05 (0.04) 0.04 (0.06) 0.04 ♂ ♂ ♂ ) (0.36) 0.39 (0.069) 0.068 (0.075) 0.1 (0.044) 0.037 (0.044) 0.075 (0.044) 0.031 (0.094) 0.062 (0.056) 0.043 (0.062 0.062 ♂ ) ( ♂ ♂ ♀ ( ( ) ( ♂ ♀ ♀ ( ♀ ( ♂ ♀ ) ♀ ♀ ♀ ( ♀ ♀ ♂ ♀ ♀ ( ♀ ♀ ♀ ♀ ♂ ♀ ♀ ( ♀ ♀ ♀ ( ♀ n. sp. n. sp. n. sp. n. sp. ♀ n. sp. sp. n. ♀ n. sp. sp. n. n. sp. n. sp. sp. n. n. sp. n. sp. n. sp. n. sp. sp. n. sp. n. n. sp. sp. n. n. sp. sp. n. n. sp. n. sp. n. sp. n. sp. n. sp. sp. n. n. sp. sp. n. n. sp. n. sp. n. sp. sp. n. n. sp. n. sp. n. Species A1L A1W A2L A2W A3L A3W A4L A4W A5L Species A2L A1L A1W T. acarinata acarinata T. acuminata T. T. acuta T. clypeata T. colombiana concavata T. T. corrugata costaricana T. delsinnei T. T. depressa T. dianae fernandezi T. T. hansoni kaulbarsi T. magnifica T. T. mexicana panamana T. T. pecki procera T. proximata T. T. punctata sculpturata T. T. transversata T. triangulata

NEOTROPICAL TRICHACIS Zootaxa 3337 © 2012 Magnolia Press · 47 0.04 (0.04) (0.04) 0.04 (0.07) 0.07 (0.05) 0.07 (0.07) 0.08 (0.05) 0.08 (0.07) 0.04 (0.05) 0.08 (0.07) 0.06 0.05 (0.06) (0.06) 0.05 (0.11) 0.09 (0.06) 0.07 (0.1) 0.09 (0.06) 0.07 (0.1) 0.09 (0.07) 0.07 (0.1) 0.09 0.037 0.037 0.044 0.056 0.056 0.069 0.062 0.069 0.081 0.04 0.04 0.07 0.07 0.06 0.08 0.09 0.09 0.09 0.06 0.04 0.069 0.04 0.04 0.075 0.06 0.06 0.05 0.06 0.08 0.08 0.08 0.07 0.07 0.09 0.06 0.07 0.06 0.09 0.07 0.09 0.09 0.09 0.09 0.09 0.1 0.1 0.08 0.06 0.04 0.05 0.05 0.08 0.07 0.06 0.06 0.06 0.06 0.06 0.08 0.08 0.06 0.08 0.08 0.09 0.08 0.04 (0.04) (0.04) 0.04 (0.08) 0.06 (0.05) 0.05 (0.09) 0.06 (0.05) 0.06 (0.08) 0.06 (0.06) 0.04 (0.08) 0.06 0.05 0.05 0.07 0.07 0.06 0.05 0.07 0.06 0.08 0.03 0.07 0.09 0.03 0.09 0.04 0.08 0.06 0.05 0.05 0.06 0.05 0.06 0.07 0.05 0.07 0.03 0.05 0.05 0.05 0.04 0.06 0.04 0.09 0.07 0.04 0.1 0.07 0.09 0.09 0.1 0.05 0.06 0.07 0.07 0.06 0.07 0.07 0.07 0.07 0.04 (0.04) (0.04) 0.04 (0.07) 0.05 (0.04) 0.05 (0.07) 0.05 (0.05) 0.06 (0.08) 0.08 (0.05) 0.08 (0.07) 0.06 0.04 (0.04) (0.04) 0.04 (0.06) 0.07 (0.05) 0.06 (0.07) 0.08 (0.06) 0.08 (0.07) 0.07 (0.06) 0.08 (0.07) 0.07 ) (0.05) 0.05 (0.06) 0.06 (0.06) 0.06 (0.07) 0.06 (0.05) 0.08 (0.07) 0.07 (0.04) 0.1 (0.07) 0.06 ) (0.04) 0.03 (0.07) 0.06 (0.05) 0.06 (0.08) 0.06 (0.05) 0.07 (0.08) 0.06 ) (0.06) 0.07 (0.08) 0.06 ) ) (0.05) 0.04 (0.07) 0.06 (0.06) 0.06 (0.09) 0.06 (0.07) 0.07 (0.08) 0.06 (0.07) 0.07 (0.07) 0.07 ) (0.04) 0.03 (0.06) 0.04 (0.06) 0.04 (0.06) 0.05 (0.06) 0.05 (0.06) 0.07 (0.06) 0.05 (0.06) 0.07 ♂ ♂ ♂ ) (0.05) 0.05 (0.056) 0.056 (0.056) 0.06 (0.06) 0.068 0.06 (0.075) 0.06 (0.075) (0.05) 0.089 (0.06) 0.081 ♂ ) ( ♂ ♂ ♀ ( ( ) ( ♂ ♀ ♀ ( ♀ ( ♂ ♀ ) ♀ ♀ ♀ ( ♀ ♀ ♂ ♀ ♀ ( ♀ ♀ ♀ ♀ ♂ ♀ ♀ ( ♀ ♀ ♀ ( ♀ n. sp. n. sp. n. sp. n. sp. ♀ n. sp. sp. n. ♀ n. sp. sp. n. n. sp. n. sp. sp. n. n. sp. n. sp. n. sp. n. sp. sp. n. sp. n. n. sp. sp. n. n. sp. sp. n. n. sp. n. sp. n. sp. n. sp. n. sp. sp. n. n. sp. sp. n. n. sp. n. sp. n. sp. sp. n. n. sp. n. sp. n. Species A5W A6L A6W A7L A7W A8L A8W A9L A8W A8L A7W A7L A6W A6L Species A5W T. acarinata acuminata T. T. acuta T. clypeata T. colombiana concavata T. T. corrugata costaricana T. delsinnei T. T. depressa T. dianae fernandezi T. T. hansoni kaulbarsi T. magnifica T. T. mexicana panamana T. T. pecki procera T. proximata T. T. punctata sculpturata T. T. transversata T. triangulata

48 · Zootaxa 3337 © 2012 Magnolia Press ARIAS-PENNA ET AL. 0.07 (0.05) (0.05) 0.07 (0.13) 0.1 (0.04) 0.06 na (0.056) 0.06 0 (0) (0.068) 0.07 (0.1) 0.07 (0.1) 0.1 0.07 (0.07) (0.07) 0.07 (0.18) 0.14 (0.06) 0.07 na (0.10) 0.094 (0.125) 0.12 (0.19) 0.13 (0.18) 0.17 (0.15) 0.16 0.09 0.12 0.08 na 0.1 0.09 0.03 0.07 0.1 0.1 0.12 0.07 0.22 0.1 0.03 0.06 0.15 0.12 0.09 0.1 0.068 0.12 0.075 0.094 0.025 0 0.1 0.07 na 0 0.08 0.075 na 0.12 0.09 0.1 0.06 na 0.1 0.07 0.19 0.075 na 0.125 0.09 0.07 0.15 0.09 0.09 0.075 0.16 0.16 0.09 0.15 na 0.16 0.16 0 0 0 0.13 0.1 0 0.04 na 0.08 0.14 0.09 na na 0.06 0.08 0.13 0.06 0.015 0.09 0.14 0.19 0.07 (0.06) (0.06) 0.07 (0.18) 0.13 (0.06) 0.07 na (0.06) 0.04 0 (0) (0.093) 0.069 (0.17) 0.16 (0.16) 0.14 0.08 0.11 0.07 na 0.08 0.1 0.09 0.1 0.17 0.17 0.1 0.093 0.09 0.09 0.1 0.1 0.1 0.08 0.08 0.09 0.043 na 0.08 0.04 0.14 0.07 0 0.11 0.08 0.09 na 0.15 0.068 0.2 0.06 0.06 0.016 0.06 0.07 na 0.13 0.07 0 0.1 0.07 0.05 0.07 0 0.04 na na 0.08 na 0.1 0.1 0.09 0.14 0.09 0.19 0.06 0.12 0.07 0.08 (0.05) (0.05) 0.08 (0.1) 0.11 (0.05) 0.06 na (0.11) 0.08 0 (0) (0.06) 0.05 (0.09) 0.11 (0.09) 0.12 0.08 (0.06) (0.06) 0.08 (0.14) 0.09 (0.06) 0.08 na (0.069) 0.08 (0.056) 0.05 (0.12) 0.15 (0.14) 0.16 (0.07) 0.03 ) (0.05) 0.1 (0.14) 0.11 (0.05) 0.07 na (0.03) 0.025 0 (0) (0.06) 0.06 (0.15) 0.12 (0.15) 0.18 ) (0.06) 0.07 (0.15) 0.12 (0.05) 0.06 na (0.069) 0.062 (0.11) 0.093 (0.11) 0.11 ) (0.13) 0.11 (0.1) 0.08 ) ) (0.06) 0.07 (0.14) 0.11 (0.05) 0.06 na (0.03) 0.025 0 (0) (0.11) 0.04 (0.17) 0.1 (0.18) 0.12 ) (0.05) 0.04 (0.11) 0.09 (0.05) 0.04 na (0.025) 0.19 0 (0.094) 0.05 0.1 (0.1) (0.12) 0.12 ♂ ♂ ♂ ) (0.06) 0.081 (0.13) 0.1 (0.056) 0.068 na (0.0125) 0.0125 0(0) (0.087) 0.093 (0.13) 0.08 (0.19) 0.16 ♂ ) ( ♂ ♂ ♀ ( ( ) ( ♂ ♀ ♀ ( ♀ ( ♂ ♀ ) ♀ ♀ ♀ ( ♀ ♀ ♂ ♀ ♀ ( ♀ ♀ ♀ ♀ ♂ ♀ ♀ ( ♀ ♀ ♀ ( ♀ n. sp. n. sp. n. sp. n. sp. ♀ n. sp. sp. n. ♀ n. sp. sp. n. n. sp. n. sp. sp. n. n. sp. n. sp. n. sp. n. sp. sp. n. sp. n. n. sp. sp. n. n. sp. sp. n. n. sp. n. sp. n. sp. n. sp. n. sp. sp. n. n. sp. sp. n. n. sp. n. sp. n. sp. sp. n. n. sp. n. sp. n. Species A9L A10L A10W POOCL POHOCL OHOCL AALL PLL CPAL CPAL PLL AALL OHOCL POOCL POHOCL A10W A10L Species A9L T. acarinata acarinata T. acuminata T. T. acuta T. clypeata T. colombiana concavata T. T. corrugata costaricana T. delsinnei T. T. depressa T. dianae fernandezi T. T. hansoni kaulbarsi T. magnifica T. T. mexicana panamana T. T. pecki procera T. proximata T. T. punctata sculpturata T. T. transversata T. triangulata

NEOTROPICAL TRICHACIS Zootaxa 3337 © 2012 Magnolia Press · 49 0.21 0.11 0.26 2.15 0.52 4.13 0.16 0.21 0.52 2.15 0.26 4.13 0.11 0.21 0.22 0.57 1.42 0.25 2.49 0.12 0.33 0.23 0.2 0.5 1.67 0.32 0.04 3.34 0.26 0.3 0.24 0.18 0.2 (0.19) (0.19) 0.2 (1.02) 1.09 (0.42) 0.4 (2.42) 2.72 0.3 (0.3) (0.17) 0.2 (0.17) 0.2 0.26 (0.3) (0.3) 0.26 (0.13) 0.12 (0.34) 0.34 (1.43) 1.71 (0.6) 0.52 (2.38) 3.2 (0.38) 0.21 (0.26) 0.22 (0.2) 0.21 0.22 (0.25) (0.25) 0.22 (0.15) 0.17 (0.34) 0.32 (1.23) 1.22 (0.61) 0.6 (2.01) 2.03 (0.4) 0.38 (0.25) 0.25 (0.23) 0.25 0.19 1 1.12 0.52 2.15 0.33 0.18 0.18 2.15 0.52 0.33 1.12 0.18 1 0.19 2.30 0.2 0.25 0.52 0.3 1.2 0.44 0.71 0.18 0.32 1.61 0.13 0.2 0.1 0.11 0.23 0.025 0.28 0.55 1.38 0.29 0.08 2.50 0.16 0.34 0.21 0.24 0.6 1.06 0.18 0.07 1.76 0.37 1.47 0.17 0.169 0.26 3.97 0.02 0.47 0.17 0.1 0.187 0.23 0.125 0.21 0.1 0.41 1.03 0.22 2.51 0.25 0.16 1.6 0.2 0.65 0.2 0.37 1.04 0.029 0.24 0.34 0.18 2.28 0.32 0.38 0.71 0.19 1.86 0.6 0.05 1.37 0.2 0.28 0.09 0.3 0.17 0.3 0.13 0.18 0.25 0.52 1.27 0.28 2.44 0.14 0.22 0.18 (0.19) (0.19) 0.18 (0.10) 0.07 (0.28) 0.27 (1.0) 1.02 (0.47) 0.45 (2.12) 2.26 (0.26) 0.27 (0.16) 0.15 (0.19) 0.17 0.2 (0.19) (0.19) 0.2 (0.06) 0.18 (0.31) 0.33 (1.13) 1.54 (0.5) 0.5 (2.26) 3.08 (0.3) 0.35 (0.15) 0.19 (0.18) 0.19 ) (0.21) 0.22 (0.1) 0.07 (0.32) 0.24 (1.45) 1.23 (0.45) 0.55 (3.22) 2.23 0.3 (0.32) (0.22) 0.2 (0.19) 0.23 ) (0.21) 0.19 (0.18) 0.14 (0.32) 0.27 (1.09) 1.08 (0.49) 0.41 (2.2) 2.6 (0.3) 0.35 (0.17) 0.18 (0.17) 0.18 ) ) ) (0.24) 0.17 (009) 0.05 (0.3) 0.22 (1.11) 1.44 (0.58) 0.47 (1.91) 3.06 (0.32) 0.25 (0.2) 0.13 (0.27) 0.2 ) (0.2) 0.2 (0.04) 0.07 (0.23) 0.24 (0.82) 0.91 (0.49) 0.46 (1.67) 1.97 (0.27) 0.25 (0.11) 0.12 (0.19) 0.15 ♂ ♂ ♂ ) (0.21) 0.025 (0.12) 0.13 (0.28) 0.29 (1.05) 1.23 (0.54) 0.48 (1.94) 2.56 (0.32) 0.32 (0.22) 0.21 (0.21) 0.21 ♂ ) ( ♂ ♂ ♀ ( ( ) ( ♂ ♀ ♀ ( ♀ ( ♂ ♀ ) ♀ ♀ ♀ ( ♀ ♀ ♂ ♀ ♀ ( ♀ ♀ ♀ ♀ ♂ ♀ ♀ ( ♀ ♀ ♀ ( ♀ n. sp. n. sp. n. sp. n. sp. ♀ n. sp. sp. n. ♀ n. sp. sp. n. n. sp. n. sp. sp. n. n. sp. n. sp. n. sp. n. sp. sp. n. sp. n. n. sp. sp. n. n. sp. sp. n. n. sp. n. sp. n. sp. n. sp. n. sp. sp. n. n. sp. sp. n. n. sp. n. sp. n. sp. sp. n. n. sp. n. sp. n. Species Species T1L MTSH T1W MTSL/MTSW MTSW MTSL PL2L MCL CPAW T. acarinata acarinata T. acuminata T. T. acuta T. clypeata T. colombiana concavata T. T. corrugata costaricana T. delsinnei T. T. depressa T. dianae fernandezi T. T. hansoni kaulbarsi T. magnifica T. T. mexicana panamana T. T. pecki procera T. proximata T. T. punctata sculpturata T. T. transversata T. triangulata

50 · Zootaxa 3337 © 2012 Magnolia Press ARIAS-PENNA ET AL. 0.6 (0.67) (0.67) 0.6 (0.61) 0.6 (0.075) 0.075 (0.53) 0.54 (0.087) 0.075 (0.53) 0.48 (0.05) 0.09 (0.42) 0.36 0.68 0.52 0.06 0.5 0.06 0.45 0.07 0.36 0.36 0.07 0.45 0.06 0.5 0.06 0.52 0.68 0.5 0.2 0.37 0.21 0.35 0.21 0.26 0.25 0.38 0.05 0.36 0.05 0.38 0.46 0.1 0.65 0.05 0.34 0.28 0.09 0.61 0.1 0.054 0.053 0.75 0.07 0.52 0.08 0.48 0.08 0.42 0.31 0.73 (0.68) (0.68) 0.73 (0.5) 0.5 (0.07) 0.11 (0.48) 0.47 (0.07) 0.12 (0.42) 0.43 (0.06) 0.14 (0.037) 0.33 0.56 (0.56) (0.56) 0.56 (0.47) 0.45 (0.07) 0.075 (0.46) 0.39 (0.07) 0.068 (0.41) 0.35 (0.06) 0.068 (0.35) 0.28 0.95 0.95 0.19 0.52 0.19 0.48 0.29 0.36 0.38 0.21 0.07 0.47 0.07 0.52 0.07 0.5 0.63 0.4 0.05 0.44 0.05 0.39 0.05 0.34 0.72 0.33 0.12 0.55 0.1 0.51 0.52 0.6 0.1 0.1 0.45 0.35 0.075 0.56 0.47 0.075 0.48 0.59 0.06 0.41 0.07 0.41 0.06 0.37 0.32 0.1 0.38 0.67 0.6 0.1 0.43 0.1 0.11 0.55 0.5 0.1 0.78 0.09 0.47 0.1 0.57 0.39 0.32 0.65 0.09 0.53 0.1 0.47 0.37 0.6 (0.57) (0.57) 0.6 (0.42) 0.4 (0.06) 0.06 (0.4) 0.4 (0.06) 0.08 (0.36) 0.3 (0.06) 0.06 (0.32) 0.3 0.76 (0.8) (0.8) 0.76 (0.6) 0.52 (0.08) 0.12 (0.55) 0.51 (0.1) 0.12 (0.5) 0.4 (0.07) 0.32 (0.41) 0.12 ) (0.6) 0.68 (0.45) 0.55 (0.08) 0.08 (0.45) 0.48 (0.07) 0.09 (0.38) 0.41 (0.06) 0.09 (0.32) 0.34 ) (0.66) 0.6 (0.49) 0.41 (0.069) 0.08 (0.44) 0.4 (0.06) 0.07 (0.4) 0.35 (0.06) 0.06 ) (0.33) 0.29 ) ) (0.62) 0.5 (0.58) 0.47 (0.08) 0.6 (0.58) 0.41 (0.05) 0.06 (0.51) 0.38 (0.05) 0.05 (0.47) 0.28 ) (0.52) 0.53 (0.49) 0.46 (0.04) 0.05 (0.42) 0.42 (0.04) 0.05 (0.38) 0.37 (0.04) 0.06 (0.31) 0.27 ♂ ♂ ♂ ) (0.55) 0.62 (0.46) 0.48 (0.075) 0.11 (0.54) 0.41 (0.069) 0.07 (0.47) 0.37 (0.062) 0.08 (0.36) 0.37 ♂ ) ( ♂ ♂ ♀ ( ( ) ( ♂ ♀ ♀ ( ♀ ( ♂ ♀ ) ♀ ♀ ♀ ( ♀ ♀ ♂ ♀ ♀ ( ♀ ♀ ♀ ♀ ♂ ♀ ♀ ( ♀ ♀ ♀ ( ♀ n. sp. n. sp. n. sp. n. sp. ♀ n. sp. sp. n. ♀ n. sp. sp. n. n. sp. n. sp. sp. n. n. sp. n. sp. n. sp. n. sp. sp. n. sp. n. n. sp. sp. n. n. sp. sp. n. n. sp. n. sp. n. sp. n. sp. n. sp. sp. n. n. sp. sp. n. n. sp. n. sp. n. sp. sp. n. n. sp. n. sp. n. Species T2L T2W T3L T3W T4L T4W T5L T5W T. acarinata acarinata T. acuminata T. T. acuta T. clypeata T. colombiana concavata T. T. corrugata costaricana T. delsinnei T. T. depressa T. dianae fernandezi T. T. hansoni kaulbarsi T. magnifica T. T. mexicana panamana T. T. pecki procera T. proximata T. T. punctata sculpturata T. T. transversata T. triangulata

NEOTROPICAL TRICHACIS Zootaxa 3337 © 2012 Magnolia Press · 51 0.09 (0.06) (0.06) 0.09 (0.26) 0.2 (0.04) (0.14) 0.17 (0.07) (0.07) 0.17 (0.31) 0.21 (0.05) (0.16) 0.13 (0.062) (0.062) 0.13 (0.33) 0.26 (0.04) (0.21) 0.1 0.1 0.13 0.25 0.12 0.22 0.09 0.27 0.21 0.09 0.22 0.1 (0.05) 0.19 (0.27) (0.025) (0.1) 0.25 (0.05) (0.05) 0.25 (0.24) 0.23 (0.05) (0.16) 0.37 0.37 0.2 0.06 0.08 0.23 0.17 0.23 0.24 0.14 0.16 0.11 0.12 0.3 0.2 0.17 0.15 0.24 0.26 ) (0.05) 0.09 (0.24) 0.24 (0.37) (0.12) ) (0.025) 0.09 (0.04) 0.12 (0.05) (0.09) ) ) ) (0.06) 0.1 (0.33) 0.21 (0.05) (0.17) ) (0.031) 0.1 (0.39) 0.16 (0.04) (0.14) ♂ ♂ ♂ ) 0.14 (0.056) 0.22 (0.29) (0.025) (0.13) ♂ ) ( ♂ ♂ ♀ ( ( ) ( ♂ ♀ ♀ ( ♀ ( ♂ ♀ ) ♀ ♀ ♀ ( ♀ ♀ ♂ ♀ ♀ ( ♀ ♀ ♀ ♀ ♂ ♀ ♀ ( ♀ ♀ ♀ ( ♀ n. sp. n. sp. n. sp. n. sp. ♀ n. sp. sp. n. ♀ n. sp. sp. n. n. sp. n. sp. sp. n. n. sp. n. sp. n. sp. n. sp. sp. n. sp. n. n. sp. sp. n. n. sp. sp. n. n. sp. n. sp. n. sp. n. sp. n. sp. sp. n. n. sp. sp. n. n. sp. n. sp. n. sp. sp. n. n. sp. n. sp. n. Species T6L T6W T7L T7W T. acarinata acarinata T. acuminata T. T. acuta T. clypeata T. colombiana concavata T. T. corrugata costaricana T. delsinnei T. T. depressa T. dianae fernandezi T. T. hansoni kaulbarsi T. magnifica T. T. mexicana panamana T. T. pecki procera T. proximata T. T. punctata sculpturata T. T. transversata T. triangulata

52 · Zootaxa 3337 © 2012 Magnolia Press ARIAS-PENNA ET AL. APPENDIX 2 URI Table matching terms and concepts used in this revision with the Hymenoptera Anatomy Ontology database.

Preferred Term in Term Concept URI HAO The antennal segment that is proximal to the pedicel and is connected with the head via A1 the radicle. http://purl.obolibrary.org/obo/HAO_0000908 scape The antennal segment that is the second segment of the antenna and is connected proximally with the scape and distally with A2 the flagellum. http://purl.obolibrary.org/obo/HAO_0000706 pedicel The flagellomere that is located immediately third A3 distal to the second flagellomere. http://purl.obolibrary.org/obo/HAO_0001895 flagellomere The flagellomere that is located immediately fourth A4 distal to the third flagellomere. http://purl.obolibrary.org/obo/HAO_0001896 flagellomere The flagellomere that is located immediately fifth A5 distal to the fourth flagellomere. http://purl.obolibrary.org/obo/HAO_0001885 flagellomere The flagellomere that is located immediately sixth A6 distal to the fifth flagellomere. http://purl.obolibrary.org/obo/HAO_0001897 flagellomere The flagellomere that is located immediately seventh A7 distal to the sixth flagellomere. http://purl.obolibrary.org/obo/HAO_0001898 flagellomere The flagellomere that is located immediately eighth A8 distal to the seventh flagellomere. http://purl.obolibrary.org/obo/HAO_0001899 flagellomere The flagellomere that is located immediately ninth A9 distal to the eighth flagellomere. http://purl.obolibrary.org/obo/HAO_0001900 flagellomere The flagellomere that is located immediately tenth A10 distal to the ninth flagellomere. http://purl.obolibrary.org/obo/HAO_0001884 flagellomere The signum that is submedian and located on the anterior margin of the mesoscutum and antero- antero-admedian corresponds to the site of origin of the admedian line longitudinal flight muscle. http://purl.obolibrary.org/obo/HAO_0000128 line The anatomical cluster that is composed of antenna the scape, pedicel and flagellum. http://purl.obolibrary.org/obo/HAO_0000101 antenna The anatomical structure that is delimited by area material or immaterial anatomical entities. http://purl.obolibrary.org/obo/HAO_0000146 area The anatomical cluster that is composed of the whole organism but which excludes the body antennae, legs and wings. http://purl.obolibrary.org/obo/HAO_0000182 body carina The process that is linear and external. http://purl.obolibrary.org/obo/HAO_0000188 carina The anatomical cluster that is composed of apical flagellomeres bearing multiporous clava plates in female organism. http://purl.obolibrary.org/obo/HAO_0000203 clava The margin that delimits the clypeus distally ventral (ventrally or anteriorly depending on the clypeal clypeal margin orientation of the head). http://purl.obolibrary.org/obo/HAO_0001767 margin The area that corresponds to the site of origin clypeus of the clypeo-epipharyngeal muscle. http://purl.obolibrary.org/obo/HAO_0000212 clypeus The leg segment that is connected to the body and to the trochanter via conjunctivae coxa and muscles. http://purl.obolibrary.org/obo/HAO_0000228 coxa ……continue on next page

NEOTROPICAL TRICHACIS Zootaxa 3337 © 2012 Magnolia Press · 53 APPENDIX 2. (Continued) Preferred Term in Term Concept URI HAO The compound organ that is composed of eye ommatidia. http://purl.obolibrary.org/obo/HAO_0000217 eye fore wing The wing that is located on the mesothorax. http://purl.obolibrary.org/obo/HAO_0000351 fore wing The area that is located dorsally of the ventral margin of the toruli and ventrally of the anterior ocellus medial to the inner frons margins of the eye and malar line. http://purl.obolibrary.org/obo/HAO_0001044 frons The area that is delimited by the intersection of the interorbital plane, the margin of the compound eye, the margin of the oral foramen, the occipital carina and the malar gena sulcus. http://purl.obolibrary.org/obo/HAO_0000371 gena The tagma that is located anterior to the head thorax. http://purl.obolibrary.org/obo/HAO_0000397 head hyperoccipital The carina that extends on the vertex hyperoccipi- carina between the outer orbits. http://purl.obolibrary.org/obo/HAO_0000406 tal carina median area inter-notaular The area that is located between the notauli. of the area http://purl.obolibrary.org/obo/HAO_0000520 mesoscutum interantennal The area that is elevated and extends in interantennal process between the toruli. http://purl.obolibrary.org/obo/HAO_0000422 process The area that is limited posteriorly by the posterior ocellar line and laterally by the ocellar interocellar area lateral ocellar line. http://purl.obolibrary.org/obo/HAO_0000430 triangle The interocular distance that is the shortest interorbital IOS between the inner margin of compound eyes. http://purl.obolibrary.org/obo/HAO_0000432 distance lateral area The area that is located between the notaulus lateral notaular of the and the parascutal carina. area http://purl.obolibrary.org/obo/HAO_0000466 mesoscutum The anatomical cluster that is composed of the coxa and all distal leg segments and is leg connected to the pectus. http://purl.obolibrary.org/obo/HAO_0000494 leg The anatomical line that is the shortest lateral between the margins of the median and ocellar LOL lateral ocelli. http://purl.obolibrary.org/obo/HAO_0000480 length The sclerite that is connected to the cranium along the anterior margin of the oral foramen via the anterior and posterior cranio- mandible mandibular articulations. http://purl.obolibrary.org/obo/HAO_0000506 mandible The line that delimits the periphery of an margin area. http://purl.obolibrary.org/obo/HAO_0000510 margin mesopleural The carina that crosses the mesopleuron and mesopleural carina limits ventrally the femoral depression. http://purl.obolibrary.org/obo/HAO_0000559 carina The area that is located laterally of the mesopleuron mesodiscrimen. http://purl.obolibrary.org/obo/HAO_0000566 mesopleuron The scutellum that is located on the mesoscu- mesoscutellum mesonotum. http://purl.obolibrary.org/obo/HAO_0000574 tellum ……continue on next page

54 · Zootaxa 3337 © 2012 Magnolia Press ARIAS-PENNA ET AL. APPENDIX 2. (Continued) Preferred Term in Term Concept URI HAO The anatomical cluster that is composed of the prothorax, mesothorax and the mesosoma metapectal-propodeal complex. http://purl.obolibrary.org/obo/HAO_0000576 mesosoma The tagma that is connected anteriorly to the metapectal-propodeal complex at the propodeal foramen and consists of metasoma abdominal segments. http://purl.obolibrary.org/obo/HAO_0000626 metasoma The line that extends submedially along the mesoscutum and corresponds to the median border of the site of origin of the first notaulus mesopleuro-mesonotal muscle. http://purl.obolibrary.org/obo/HAO_0000647 notaulus The multi-tissue structure that is located on the top of the head, composed of the corneal lens, pigment cell, rhabdoms and synaptic ocellus, ocelli plexus. http://purl.obolibrary.org/obo/HAO_0000661 ocellus The anatomical line that is shortest and connects the compound eye and the lateral ocular OOL ocellus. http://purl.obolibrary.org/obo/HAO_0000662 ocellar line The signum that is located between the notaulus and the parascutal carina and corresponds to the site of origin of the parapsidal parapsidal line dorsoventral indirect flight muscle. http://purl.obolibrary.org/obo/HAO_0000694 signum posterior ocellus, lateral posterior ocelli The ocellus that is paired. http://purl.obolibrary.org/obo/HAO_0000481 ocellus process The area on the sclerite that is raised. http://purl.obolibrary.org/obo/HAO_0000822 process projection The process that is located on an edge. http://purl.obolibrary.org/obo/HAO_0000829 projection pronotum The notum that is located in the prothorax. http://purl.obolibrary.org/obo/HAO_0000853 pronotum The anatomical structure that is delimited by region material or immaterial anatomical entities. http://purl.obolibrary.org/obo/HAO_0000146 area The area that is located on the sclerite and that is composed of repetitive anatomical sculpture structures. http://purl.obolibrary.org/obo/HAO_0000913 sculpture specialized new label area of The setifeous patch that is located and class for mesoscutellum posteromedially on the mesoscutellum. http://purl.obolibrary.org/obo/HAO_0001967 HAO The tergum that is located on abdominal abdominal T1 segment 2. http://purl.obolibrary.org/obo/HAO_0000053 tergum 2 The tergum that is located on the abdominal abdominal T2 segment 3. http://purl.obolibrary.org/obo/HAO_0000056 tergum 3 The tergum that is located on abdominal abdominal T3 segment 4. http://purl.obolibrary.org/obo/HAO_0000057 tergum 4 The tergum that is located on abdominal abdominal T4 segment 5. http://purl.obolibrary.org/obo/HAO_0000058 tergum 5 The tergum that is located on abdominal abdominal T5 segment 6. http://purl.obolibrary.org/obo/HAO_0000059 tergum 6 The tergum that is located on abdominal abdominal T6 segment 7. http://purl.obolibrary.org/obo/HAO_0000060 tergum 7 ……continue on next page

NEOTROPICAL TRICHACIS Zootaxa 3337 © 2012 Magnolia Press · 55 APPENDIX 2. (Continued) Preferred Term in Term Concept URI HAO The tergum that is located on abdominal abdominal T7 segment 8. http://purl.obolibrary.org/obo/HAO_0000061 tergum 8 The area that is located on the gena, posterior temple to the compound eye. http://purl.obolibrary.org/obo/HAO_0000995 temple The foramen that is located on the head in antennal torulus which the radicle is positioned. http://purl.obolibrary.org/obo/HAO_0001022 foramen The area that is delimited by the intersection of the margin of the compound eyes, the interorbital plane, and the anatomical line that is tangential to the point on the margin of the anterior ocellus which defines the minimum distance between the anterior vertex ocellus and the oral foramen. http://purl.obolibrary.org/obo/HAO_0001077 vertex

56 · Zootaxa 3337 © 2012 Magnolia Press ARIAS-PENNA ET AL.