A González-Moreno & S Bordera

Review of Mexican Species of Podogaster Brullé (Hymenoptera: Ichneumonidae: Anomaloninae) with Description of Two New Species A González-Moreno & S Bordera Neotropical Entomology ISSN 1519-566X Volume 42 Number 1 Neotrop Entomol (2013) 42:39-51 DOI 10.1007/s13744-012-0082-4 1 23

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Neotrop Entomol (2013) 42:39 51 DOI 10.1007/s13744-012-0082-4 SYSTEMATICS, MORPHOLOGY AND PHYSIOLOGY Review of Mexican Species of Podogaster Brullé (Hymenoptera: Ichneumonidae: Anomaloninae) with Description of Two New Species AGONZÁLEZ-MORENO, SBORDERA Instituto de Investigación CIBIO (Centro Iberoamericano de Biodiversidad), Univ de Alicante, Alicante, Spain Keywords Biosphere reserve, neotropics, parasitoids, taxonomic key Correspondence A González-Moreno, Instituto de Investigación CIBIO (Centro Iberoamericano de Biodiversidad), Univ de Alicante, Apdo. Corr. 99, 03080 Alicante, Spain; jandra28@yahoo.com Edited by Roberto A Zucchi ESALQ/USP Received 10 May 2012 and accepted 14 August 2012 Published online 7 September 2012 Abstract Two new species of Podogaster Brullé, Podogaster brunneus n. sp. and Podogaster lagartensis n. sp., are described. The material was collected with Malaise traps operated for a year in the Ría Lagartos Biosphere Reserve, a dry tropical area of Southeast Mexico. Podogaster rosteri Gauld & Bradshaw is synonymized with Podogaster mexicanus (Cresson). A key to the Mexican species is also provided. * Sociedade Entomológica do Brasil 2012 Introduction Podogaster Brullé is a monophyletic Neotropical genus of Anomaloninae, characterized by having a transverse flexible suture in front of the scuto-scutellar groove, fore wing with Cu1a and 1m-cu separated basally by a distance of about 0.1 the length of Cu1b or less (sometimes contiguous and rarely basally united), a long antennal pedicel, a long ovipositor, slightly hairy eyes (Gauld & Bradshaw 1997), and specialized male genitalia (Gauld 1976). Reported hosts of Podogaster (all Lepidoptera) are species of Saturniidae: Automeris io (Fabricius), Hylesia lineata Druce (Gauld & Bradshaw 1997), and Hylesia metabus (Cramer) (Hernandez et al 2009), and Pyralidae: Cactoblastis cactorum (Berg) (Cushman 1927) and Hypsipyla grandella (Zeller) (Graf 1983). The genus comprises 26 described species of exclusively Neotropical distribution. There is no complete revision of the species, but a good contribution to the genus was provided by Gauld & Bradshaw (1997), who described 14 new species from Costa Rica and provided a key for 15 reported species. The species Podogaster eldae Gauld & Bradshaw, Podogaster mexicanus (Cresson), Podogaster rosteri Gauld & Bradshaw, Podogaster ruthae Gauld & Bradshaw, Podogaster vitticollis (Cresson) and Podogaster tranae Gauld & Bradshaw have been previously reported from Mexico (Cresson 1874, Cameron 1886, Gauld & Bradshaw 1997, Ruíz-Cancino et al 2002, González-Moreno & Bordera 2011). The aim of this study is to describe and illustrate two new species from Mexico and to provide a new key for the reviewed Mexican species of Podogaster. Material and Methods Part of the material was collected using Malaise traps in the dry forest of the Ría Lagartos Biosphere Reserve (Reserva de la Biósfera Ría Lagartos), approximately 21 36 N, 88 10 W in Northeastern Yucatán. Malaise trap collecting pots were replaced fortnightly from June 2008 to August 2009. Specimens of P. eldae and P. ruthae preserved at the Insect Museum of Universidad Autónoma de Tamaulipas (UAT, Ciudad Victoria, Mexico) were revised. Type material of Podogaster species reported by Gauld & Bradshaw (1997), preserved at the Instituto Nacional de

40 González-Moreno & Bordera Biodiversidad (INBio, Santo Domingo de Heredia, Costa Rica), American Entomological Institute (AEIC, Gainesville, Florida, USA), and the Natural History Museum (BMNH, London, UK), as well as the type specimens of Podogaster major Szépligeti, Podogaster medius Szépligeti, Podogaster minor Szépligeti, and Podogaster variegatus Szépligeti kept at the Termeszettudomanyi Muzeum Allattara (TMA, Budapest, Hungary) were studied. Images of the type specimens of P. mexicanus (Cresson) made by Jason Weintraub from the Academy of Natural Sciences of Philadelphia (ANSP, Pennsylvania, USA) and P. rosteri Gauld & Bradshaw made by Ronald Zuñiga from INBio (Costa Rica) were examined for comparison. The remaining species have been checked by their original descriptions. Morphological and body surface sculpturing terminology follows Gauld & Bradshaw (1997). Key has been modified also from Gauld & Bradshaw (1997). Layered digital images were made at the University of Turku (Finland), with an Olympus E520 digital camera attached to an Olympus SZX16 stereomicroscope and then combined using Deep Focus 3.1. The SEM images were taken using a Hitachi S-3000N (in low vacuum mode) at the University of Alicante, Spain. The material of new species is deposited in the Colección Entomológica Regional of Universidad Autónoma de Yucatán (CER-UADY, Mérida, Yucatán, Mexico) and in the Colección Entomológica of Universidad de Alicante (CEUA, Alicante, Spain). Results Key to Mexican species of Podogaster 1. Head in dorsal view with gena strongly swollen (Fig 3), in lateral view width of gena at midpoint almost equal to width of eye at this point (Figs 1, 7); fore wing with length of first abscissa of Cu1a 0.40 0.65 times length of Cu1 between 1m-cu and cu-a (Figs 1, 5); clypeus strongly produced ventrally (Figs 2, 6); eyes in frontal view weakly convergent ventrally (Figs 2, 6)......P. mexicanus (Cresson) (0P. rosteri Gauld & Bradshaw, n. syn.) Head in dorsal view with gena not swollen, or only slightly swollen (Figs 17, 18), clearly less than 0.5 times width of eye in lateral view (Figs 8, 10, 11, 13); fore wing with length of first abscissa of Cu1a 0.8 1.6 times length of Cu1 between 1m-cu and cu-a (Figs 8, 12, 14); clypeus slightly produced ventrally (Figs 15 and 16); eyes in frontal view strongly convergent ventrally (Figs 15, 16)... 2 2. Fore wing with length of first abscissa of Cu1a 1.12 1. 59 times length of Cu1 between 1m-cu and cu-a, without a pigmented region resembling a vein extending from lower distal corner of first subdiscal cell to hind margin of wing (Fig 8, arrow a), apex hyaline or very weakly infumate; antenna with 24 to 32 flagellomeres. 3 Fore wing with length of first abscissa of Cu1a 0.8 1. 20 times length of Cu1 between 1m-cu and cu-a, with a pigmented region resembling a vein extending from lower distal corner of first subdiscal cell to hind margin of wing (Fig 12, arrow a), apex usually strongly infumate (Figs 12, 14); antenna with 35 to 51 flagellomeres........ 4 3. Posterior apex of propodeum extended into a slender, unsculptured neck; fore wing with 1m-cu and Cu1a arising from a short common stalk (Fig 8, arrow b). Tergite I of metasoma with spiracle small, not larger than the spiracle of tergite II, and separated from hind margin of tergite by about 1.2 1.5 the dorsal interspiracular distace. Scutellum yellow with a median longitudinal black line......p. tranae Gauld & Bradshaw Posterior apex of propodeum evenly tapered into short sculptured neck; fore wing with 1m-cu and Cu1a basally separated by a short abscissa of Cu1 that at most is about 0.15 of length of Cu1b. Tergite I of metasoma with spiracle enlarged, bigger than the spiracle of tergite II, and separated from hind margin of tergite by about the dorsal interspiracular distance. Scutellum entirely yellow......p. ruthae Gauld & Bradshaw 4. Mesosoma mainly black, with yellow markings only on the pronotum, mesoscutum and scutellum, propodeum being virtually entirely black......c. eldae Gauld & Bradshaw Mesosoma mainly yellow and/or brown, sometimes with black marks on mesoscutum, propodeum, and scarcely on mesopleuron (Figs 9 11, 13).... 5 5. Scutellum coarsely rugose, broadly concave centrally; pronotum laterally and mesopleuron longitudinally striate, with only a smooth area just below the speculum (Fig 9). P. vitticollis (Cresson) Scutellum smooth and shiny, very slightly concave centrally (Figs 17 19, 21); pronotum laterally smooth and shiny; mesopleuron very slightly striate on upper part, median area at level of speculum smooth, lower part punctate and weakly striate on anterior margin (Figs 20, 22)..... 6

Mexican Species of Podogaster 41 Figs 1-4 Podogaster mexicanus, lectotype, : 1 head and mesosoma, lateral view; 2 head, frontal view; 3 head and mesoscutum, dorsal view; 4 metasoma, lateral view. 6. Mesoscutum finely punctate on a granulate background on median and lateral lobes, rugose in notaular and internotaular posterior areas (Figs 17, 19); mesopleuron finely punctate on lower part (Fig 20); propodeum with pubescence relatively short, sculpture clearly visible (Figs 10, 11, 20); pterostigma and veins dark brown in dorsal view, yellowish to light brown ventrally (Fig 12); eyes and ocelli light gray (Figs 15, 17)...... P. lagartensis n. sp. Mesoscutum strongly and densely punctate on a slightly granulate background, strongly rugose in notaular and internotaular posterior areas (Figs 18, 21); mesopleuron coarsely punctate on lower part (Fig 22); propodeum with pubescence relatively long, partially hiding sculpture (Figs 13, 22); pterostigma and veins dark brown dorsally and ventrally (Fig 14); eyes and ocelli dark brown to black (Figs 16, 18).. P. brunneus n. sp. Podogaster brunneus González-Moreno & Bordera n. sp. (Figs 13, 14, 16, 18, 21 23) Type material (11 )

42 González-Moreno & Bordera Figs 5 7 Podogaster rosteri, holotype, : 5 habitus, lateral view; 6 head, frontal view; 7 head and mesoscutum, dorsal view. Holotype (CEUA): Mexico, Yucatán State, Ría Lagartos Reserve, Cuyo Station, Sacbo, dry forest, 27.VII 6.VIII.2009, A. González-Moreno leg., Malaise trap., dry mounted. Paratypes: 1 (CER-UADY): same locality, 28.VIII 10.IX.2008; 1 (CEUA): same locality, 26.XI 10.XII.2008; 1 (CEUA): same locality, 08 22.VII.2009; 1 (CER-UADY): Tekal, dry forest, 17.VI 1.VII.2008; 1 (CEUA): same locality, 12 28.VIII.2008; 3 (CER-UADY): same locality, 28.VIII 10.IX.2008; 1 (CEUA): same locality, 4 18.II.2009; 1 (CEUA): Ría Lagartos Station, Core II, Savannah, 22.X 10.XI.2008. All of them A. González-Moreno leg., Malaise trap, dry mounted. Diagnosis As P. lagartensis, this species can be distinguished from all other known Neotropical species mainly because it is a yellowish orange species with very few black marks (only the interocellar area and sometimes the occiput). Podogaster brunneus belongs to the P. vitticollis-species group of Gauld & Bradshaw (1997) in having head in dorsal view with gena not swollen, less than 0.5 times width of eye in lateral view; and fore wing with first abscissa of Cu1a 0.9 1.0 times as long as Cu1 between 1m-cu and cu-a; a pigmented region resembling a vein extending from lower distal corner of first subdiscal cell to hind margin of wing; apex usually strongly infumate; antenna with 38 to 49 flagellomeres; clypeus slightly produced and eyes strongly convergent ventrally. P. brunneus differs from P. eldae and P. vitticollis by having the mesosoma mostly brownish, sometimes with slightly black marks on mesoscutum; and the scutellum smooth, shiny and very slightly concave centrally; pronotum laterally smooth and shiny; mesopleuron very slightly striate on upper part, median area at level of speculum smooth and lower part of mesopleuron punctate and striate on anterior margin. Podogaster brunneus also

Mexican Species of Podogaster 43 Figs 8-14 Podogaster spp.: 8 Podogaster tranae,, habitus, lateral view (arrows a area without a pigmented region resembling a vein; b 1m-cu and Cu1a arising from a short common stalk); 9 Podogaster vitticollis,, mesosoma, lateral view; 10 12 Podogaster lagartensis, 10 habitus lateral, 11 habitus lateral, 12 wings (arrow a pigmented region resembling a vein), 13 14 Podogaster brunneus, 13 habitus lateral, 14 wings. differs from P. lagartensis in having the mesoscutum strongly and densely punctate on a slightly granulate background, strongly rugose in notaular and posterior internotaular areas, median and lateral lobes slightly granulate; mesopleuron coarsely punctate on lower part; propodeum strongly pubescent with hairs relatively long and partially hiding sculpture; pterostigma and veins dark brown dorsally and ventrally; and eyes and ocelli dark brown to black.

44 González-Moreno & Bordera Figs 15-18 Podogaster spp.: 15, 16 Head, frontal view, 15 Podogaster lagartensis, 16 Podogaster brunneus, 17, 18 Head and anterior part of mesosoma, dorsal view, 17 Podogaster lagartensis, 18 Podogaster brunneus. Description FemaleR Body length (without antennae and ovipositor) 8 15 mm; fore wing 3.9 7.3 mm (holotype: body length 14.8 mm; fore wing 7.2 mm). HeadR Lower face finely punctate, strongly narrowed ventrally, its width at level of clypeal suture about 0.4 times width at level of base of antennal sockets; clypeus slightly produced, with a median apical strong tooth (Fig 16); mandibles slender, upper tooth slightly longer than lower tooth; lower margin of eye contiguous with base of mandible (Fig 16), eyes strongly convergent ventrally; distance between lateral ocellus and eye 0.9 1.0 times maximum diameter of ocellus (Fig 18); frons with moderately strong punctures, without a median vertical carina; gena in dorsal view not swollen, less than 0.5 times

Mexican Species of Podogaster 45 Figs 19-22 Podogaster spp., mesosoma: 19, 20 Podogaster lagartensis, 19 dorsal view, 20 lateral view, 21, 22 Podogaster brunneus, 21 dorsal view, 22 lateral view. width of eye in lateral view; occipital carina complete (Fig 18), ventrally joining hypostomal carina above base of mandible; antenna with 38 49 flagellomeres. MesosomaR Pronotum mostly smooth, with upper and posterior margin and lower corner slightly punctate, lower posterior margin striate (Fig 13); epomia strong, not arising from a discernible tooth, weakly diverging from anterior margin of pronotum, joining upper margin of pronotum at about 90 ; pronotum laterally smooth and shiny; mesoscutum strongly and densely punctate and granulate, rugose in notaular and posterior internotaular areas, median and lateral lobes slightly granulate, median lobe rounded anteriorly; scutellum transverse, smooth and shiny, weakly concave centrally, without lateral carinae (Figs 18, 21); mesopleuron ventrally coarsely punctate, grading to striate at anterior margin, upper part and mesepimeron striate with smooth area running longitudinally from anterior margin of mesopleuron to area just below speculum; epicnemial carina strong, medioventrally forming a lobe that protrudes between anterior coxae, extending about 0.5 of way up anterior pleural margin, moderately sinuous in front of sternaulus; sternaulus present in the form Figs 23-24 Phenology: 23 Podogaster brunneus, 24 Podogaster lagartensis.

46 González-Moreno & Bordera of a row of shallow foveae extending across about 0.5 of mesopleuron; sternal region punctate. Propodeum and metapleuron reticulately sculptured, with long white hairs; propodeal apex extended into a short reticulate neck (Figs 13, 22). Fore wing with a pigmented region resembling a vein extending from lower distal corner of first subdiscal cell to hind margin of wing, 1m-cu and Cu1 basally often separated by a short abscissa of Cu1, but always with this abscissa about 0.1 0.2 times length of Cu1b; length of first abscissa of Cu1a 0.9 1.0 times length of Cu1 between 1m-cu and cu-a; hind wing with posterolateral corner of sub-basal cell about 100 ; distal abscissa of 1A and Rs present as stub (Figs 13, 14). MetasomaR Slender, elongate, strongly compressed; posterior 0.3 of tergite I swollen, spiracle small, separated from hind margin of tergite by 1.0 times the interspiracular distance; ovipositor 0.9 1.1 times length of hind tibia (Fig 13). ColorR Yellow-brown to orange. Head yellow with interocellar area black; occiput orange brown to black; eyes and ocelli dark brown to black (Figs 16, 18); antenna changing gradually from orange basally to black apically, with scape yellow, pedicel ventrally yellow, dorsally orange. Mesosoma orange, with upper posterior corner of pronotum, notaular area and scutellum yellow. Metasoma orange, tergite II fuscous dorsally. Fore and mid legs yellow, mid tarsus fuscous at apex; hind leg orange with coxa dorsally yellow and tarsus infuscate. Wings hyaline, tinged with fuscous at apex, pterostigma and veins dark brown in dorsal and ventral view. Ovipositor sheath black. VariationR Some specimens have a central or three dark longitudinal patches on the mesoscutum, especially small specimens. MaleR Unknown. HostsR Unknown. DistributionR Mexico (Yucatán). HabitatR Specimens were collected in a dry forest and savannah, the latter characterized by dwarf vegetation with a low density of trees, which usually belong to one or two species. This vegetation is able to endure long inundation periods as well as several months of drought. The most common species is Plumeria obtusa (Apocynaceae). EtymologyR This species is named after the predominantly brownish colour of the body. PhenologyR See Fig 23. Podogaster eldae Gauld & Bradshaw 1997 Podogaster eldae Gauld & Bradshaw 1997: 137. Holotype. Type material (4 &1 ) Holotype (INBio): Costa Rica, Guanacuaste National Park, Maritza Station, Volcan Orosi, 560 m, III IV.1990, Gauld & Mitchell leg. Partypes: 1 (INBio): Costa Rica, Heredia Prov., La Selva Biological Station, 50 150 m, 10 26 N, 84 01 W, primary forest, 2.V.1993, INBio-OET, M/04/099; 1 (INBio): same locality and vegetation, 10 26 N, 84 01 W, 18.V.1993, INBio-OET, M/04/099; 1 (INBio): same locality, Finca La Selva, 3 km S Sarapiquí, 50 a 100 m, 1 31.III.1993, unknown leg., INBio-OET, L_N_268800_535300 #44900; 1 (INBio): same locality, 1 31.X.1993, unknown leg., manual, INBio-OET, L_N_268000_535600 #62677. Non-type material (2 &1 ) 1 (UAT): Mexico, Tamaulipas, Villa Casas Soto la Marina, Km 68 Victoria. 3.I.1987, E. Ruíz C. leg.; 1 (UAT): same locality and collector, 18.IV.1987; 1 (UAT): G. Farias, Cedros, N. FOR. YPT. 94037, 18 19.VII.1994, J.B. Woolley leg. Diagnosis Podogaster eldae differs from all other Mexican species by the following combination of characters: gena not swollen, less than 0.5 times width of eye in lateral view; fore wing with first abscissa of Cu1a 0.8 1.2 times as long as Cu1 between 1m-cu and cu-a, with a pigmented region resembling a vein extending from lower distal corner of first subdiscal cell to hind margin of wing, apex usually strongly infumate; antenna with 36 to 51 flagellomeres; clypeus slightly produced ventrally; eyes in fontal view strongly convergent ventrally; mesosoma mainly black, with yellow markings only on the pronotum, mesoscutum, and scutellum, propodeum being virtually entirely black. VariationR In the material from Mexico, females have the propodeum mainly black with two longitudinal reddish hues on both sides. HostsR Unknown. DistributionR Neotropical. Costa Rica, Guyana, Peru, Mexico (Tamaulipas). HabitatR Gauld & Bradshaw (1997) provided data on this common species in Costa Rica, widely collected from sea level up to about 1,500 m, generally in humid

Mexican Species of Podogaster 47 forest. In Mexico, this has been collected in Riparian forest (Ruíz-Cancino 2010). PhenologyR In Costa Rica, this species has been collected abundantly throughout the year; in Guyana, in February and in Peru, from March to August (Gauld & Bradshaw 1997). In Mexico, it was collected in January, March, April (Ruíz-Cancino 2010), and July. Podogaster lagartensis González-Moreno & Bordera n. sp. (Figs 10 12, 15, 17, 19 21, 24) Type material (6 &4 ) Holotype (CEUA): Mexico, Yucatán State, Ría Lagartos Reserve, Cuyo Station, Sacbo, dry forest, 20.V 3.VI.2008, A. González-Moreno leg., Malaise trap, dry mounted. PAR- ATYPES: 3 (CEUA): same locality and date as holotype; 2 (CER-UADY): same locality, 3 17.VI.2008; 4 (CEUA): same locality, 3 17.VI.2008. All of them A. González- Moreno leg., Malaise trap, dry mounted. Diagnosis This species can be distinguished from all other known Neotropical species mainly because it is almost entirely yellowish with very few black parts (only the interocellar area and sometimes the occiput). Podogaster lagartensis belongs to the P. vitticollis-species group of Gauld & Bradshaw (1997) as the head in dorsal view has the gena not or only slightly swollen, clearly less than 0.5 times width of eye in lateral view; and fore wing with first abscissa of Cu1a 0.9 1.0 times as long as Cu1 between 1m-cu and cu-a, witha pigmented region resembling a vein extending from lower distal corner of first subdiscal cell to hind margin of wing, apex usually strongly infumate; antenna with 47 to 48 flagellomeres; clypeus slightly produced ventrally and eyes strongly convergent ventrally. P. lagartensis differs from P. eldae and P. vitticollis by having the mesosoma entirely yellowish; and scutellum smooth, shiny and very slightly concave centrally; pronotum laterally smooth and shiny; mesopleuron very slightly striate on upper part, median area at level of speculum smooth and lower part of mesopleuron punctate and striate on anterior margin. P. lagartensis is distinguishable from the closest species P. brunneus in having mesoscutum finely punctate, rugose in notaular area and in the internotaular posterior areas, granulate on median and lateral lobes; mesopleuron finely punctate on lower part; propodeum strongly pubescent, hairs relatively short, not hiding sculpture; pterostigma and veins dark brown in dorsal view, yellowish to light brown ventrally; eyes and ocelli light gray. Description FemaleR Body length (without antennae and ovipositor) 12.0 13.0 mm; fore wing 6.0 7.0 mm (Holotype: body length 12.8 mm; fore wing 6.4 mm). HeadR Lower face finely punctate, narrowed ventrally, width at clypeal suture about 0.4 times width at base of antennal sockets; clypeus slightly convex, with a median apical strong tooth (Fig 15); mandibles slender, upper tooth slightly longer than lower tooth; lower margin of eye contiguous with base of mandible (Fig 15), eyes strongly convergent ventrally; distance between lateral ocellus and eye 0.9 1.1 times maximum diameter of ocellus (Fig 17); frons finely punctate, without a median vertical carina; gena in dorsal view not swollen, clearly less than 0.5 times width of eye in lateral view; occipital carina complete (Fig 17), ventrally joining hypostomal carina above base of mandible; antenna with 47 48 flagellomeres. MesosomaR Pronotum mostly smooth with upper, posterior margin and lower corner finely punctate, lower posterior margin striate (Fig 10); epomia strong, not arising from a discernible tooth, weakly diverging from anterior margin of pronotum, joining upper margin of pronotum at about 90 ; mesoscutum finely punctate, rugose in notaular and internotaular posterior areas, granulate on median and lateral lobes; median lobe rounded anteriorly; scutellum transverse, smooth and shiny, with a few longitudinal striae on posterior central margin, weakly concave, without lateral carinae (Figs 17, 19); mesopleuron ventrally finely punctate and grading to rugose striate anteriorly, upper part and mesepimeron striate with smooth area running longitudinally from anterior margin of mesopleuron to area just below speculum; epicnemial carina strong, medioventrally forming a lobe that protrudes between anterior coxae, extending about 0.5 of way up anterior pleural margin, moderately sinuous in front of sternaulus; sternaulus weak, present in the form of a row of shallow foveae extending across about anterior third of mesopleuron; sternal region punctate. Propodeum and metapleuron reticulately sculptured, with white short hairs denser on the latter; reticulated sculpture of propodeum clearly visible; propodeal apex extended into a short reticulate neck (Figs 10, 20). Fore wing with a pigmented region resembling a vein extending from lower distal corner of first subdiscal cell to hind margin of wing; 1m-cu and Cu1 basally often separated by a short abscissa of Cu1, but always with this abscissa about 0.1 times length of Cu1b; length of first abscissa of Cu1a 0.9 1.0 times length of Cu1 between 1m-cu and cu-a; hind wing with posterolateral corner of sub-basal cell about 100 ; distal abscissa of 1A and Rs present as stub (Figs 10, 12).

48 González-Moreno & Bordera MetasomaR Slender, elongate, strongly compressed; posterior 0.3 of tergite II swollen, spiracle small, separated from hind margin of tergite by 0.8 1.1 times the interspiracular distance; ovipositor 0.9 1.1 times length of hind tibia (Fig 10). ColorR Yellow-orange. Head yellow with interocellar area black; occiput orange; eyes and ocelli bright light gray (Figs 15, 17); antenna gradually changing from orange basally to black apically, with scape entirely yellow and pedicel ventrally yellow and dorsally orange. Propleuron yellow beneath. Mesosoma orange, with upper posterior and lower corners of pronotum, notaular area, and scutellum yellow beneath. Metasoma orange, tergite II fuscous dorsally. Fore and mid legs yellow with coxae and trochanters light yellowish, mid tarsus fuscous at apex; hind leg orange with coxa yellow ventrally, orange dorsally, and tarsus infuscate. Wings hyaline, tinged with fuscous at apex, with pterostigma and veins dark brown in dorsal view, yellowish to light brow ventrally (Figs 10, 12). MaleR Body length is 12.0 mm; fore wing is 7.0 mm. Similar to female in structure and coloration (Fig 11). Antenna with 46 47 flagellomeres; claspers apically rounded. Occiput mostly black; mesopleuron more yellowish than orange; tergite II dorsally fuscous or black; hind coxa yellow with posterior orange-brown to black mark; trochanter laterally dark brown, tibia ventrally yellowish, tarsomeres fuscous. HostsR Unknown. DistributionR Mexico (Yucatán). HabitatR Specimens were collected in a dry forest characterized by trees reaching 10 20 m in height, 50 75% of which drop their leaves during the dry season, and a shrub and herbaceous layer that develops during the rainy season. Epiphytic plants are abundant (Flores & Espejel 1994). Common species in this dry forest include Acacia pennatula, Caesalpinia gaumeri, Lysiloma latisiliquum, Enterolobium cyclocarpum, Piscidia piscipula and Pithecellobium brownii (Fabaceae), Bursera simaruba (Burseraceae), Guazuma ulmifolia (Sterculiaceae), Vitex gaumeri (Verbenaceae), and Metopium brownei (Anacardiaceae). EtymologyR This species is named after the type locality, Ría Lagartos Biosphere Reserve. PhenologyR See Fig 24. Podogaster mexicanus (Cresson) (Figs 1 7) Anomalon mexicanum Cresson 1874: 376. Type. Podogaster rosteri Gauld & Bradshaw 1997: 133. Type. Syn. nov. Type material (2 ) As P. mexicanus: lectotype : Mex. [0 Mexico], 68 TYPE No. 832 (Images ANSP). As P. rosteri: holotype (INBio): Costa Rica, Guanacaste Province, Santa Rosa National Park, 250 300 m, 1984, Janzen leg. Material from Mexico reported by Gauld & Bradshaw (1997) and deposited in BMNH was not found (G. Broad pers. comm.). Diagnosis Head in dorsal view with gena strongly swollen, in lateral view, width of gena at midpoint almost equal to width of eye at this point; fore wing with first abscissa of Cu1a 0.40 0.65 times as long as Cu1 between 1m-cu and cu-a, clypeus strongly produced; eyes weakly convergent ventrally; and body mostly yellowish orange except black marks on ocellar area, frons, occiput, center of the lobes of mesoscutum, an elongate patch on the mesopleuron, anterior lateral margin of propodeum, metapleuron, part of hind coxa, trochanter, and trochantellus. HostsR A multivoltine species reported as P. rosteri, from species of Saturniidae (Lepidoptera) (Gauld & Bradshaw 1997). DistributionR Neotropical. Costa Rica, Mexico (Colima). HabitatR In Costa Rica, specimens were collected in the seasonally dry forest in North-western Guanacaste. PhenologyR September November, January March, and June (Gauld & Bradshaw 1997). RemarksR Podogaster rosteri is conspicuously similar to P. mexicanus by having head in dorsal view with gena strongly swollen; fore wing with first abscissa of Cu1a about half times length of Cu1 between1m-cu and cu-a; clypeus strongly produced ventrally; eyes weakly convergent ventrally; and mostly yellowish orange body (Figs 1, 2, 3, 4, 5, 6, and7). None relevant morphological and coloration differences have been found, with exception of the size and form of black mark on the mesopleuron of both species (Types). Podogaster mexicanus has mesopleuron mostly black except posterior and dorsal yellow margins (Fig 1), while P. rosteri has a black elongate patch, which is partially separated from black lower half of mesopleuron and sternal region (Fig 5). Nevertheless, according to Gauld and Bradshaw (1997), paratypes of P. rosteri from Mexico have more extensive black on mesopleuron covering all it except for band of yellow along the posterior and dorsal

Mexican Species of Podogaster 49 margins and this is as in the lectotype of P. mexicanus. On the other hand, in two paratypes of P. rosteri from Costa Rica preserved in BMNH, this large black mark on the mesopleuron is narrowly divided into two marks (G. Broad pers. comm.). In the absence of other diagnostic characters, we consider these coloration differences as intraspecific variability and consequently P. rosteri becomes a synonym of P. mexicanus. Thus the P. rosteri species group defined by Gauld & Bradshaw (1997) must be renamed to P. mexicanus species group. This species-group comprises now P. mexicanus from Costa Rica and Mexico, and two species from Peru and Brazil, which differ from P. mexicanus by having longer ovipositor and body mostly blackish (Gauld & Bradshaw 1997). Podogaster ruthae Gauld & Bradshaw Podogaster ruthae Gauld & Bradshaw 1997: 149. Holotype. Type Material (1 ) Holotype (INBio): Costa Rica, Cartago Prov., Turrialba, Grano de Oro on Chirripo, 1,120 m, III.1993, Campos leg. Non-type material (1 ) 1 (UAT): Mexico, Tamaulipas, G. Farias, dry forest, 13 19.XII.1990, E. Ruiz C. leg. DiagnosisR This species differs from other Mexican species by the following combination of characters: gena less than 0.5 times width of eye in lateral view; fore wing with first abscissa of Cu1a 1.26 1.40 times as long as Cu1 between 1mcu and cu-a, and without a pigmented region resembling a vein extending from lower distal corner of first subdiscal cell to hind margin of wing, apex hyaline or very weakly infumate; antenna with 29 to 30 flagellomeres; clypeus slightly produced ventrally; eyes strongly convergent ventrally; posterior apex of propodeum evenly tapered into short sculptured neck; fore wing with 1m-cu and Cu1a basally separated by a short abscissa of Cu1 that at most is about 0.15 of length of Cu1b; tergite I of metasoma with spiracle enlarged, bigger than the spiracle of tergite II, and separated from hind margin of tergite by about the dorsal interspiracular distance; scutellum entirely yellow. RemarksR The specimen from Mexico differs from original description (Gauld & Bradshaw 1997) in having the mesoscutum granulate in median and lateral lobes, being shiny in notaular area, with coarse rugae on smooth background; propodeum mainly yellow, being the black area reduced only to anterior margin; hind coxae reddish brown and dorsally yellow; and a short and very weakly pigmented vein extending from lower distal corner of first subdiscal cell to hind margin of wing. Apart from all these differences, the specimen agrees with all other characters of C. ruthae. Nevertheless we do not exclude the possibility that the specimen could belong to a new species, but the study of new material is needed to solve this question. HostsR Unknown. DistributionR Neotropical. Costa Rica, Mexico (Tamaulipas). HabitatR This species have been collected in a great variety of habitats, from suburban gardens to wet and dry forest and an altitude of 1,300 m (Gauld & Bradshaw 1997). In Mexico, this has been collected in dry forest. PhenologyR In Costa Rica, this species have been collected from March to May (Gauld & Bradshaw 1997). In Mexico, it was collected at December. Podogaster tranae Gauld & Bradshaw Podogaster tranae Gauld & Bradshaw 1997: 151. Holotype. Type material (4 &1 ) Holotype (INBio): Costa Rica, Puntarenas Prov., Golfo Dulce Forest Reserve, 24 kmw Pan American highway on road to Rincon de la Osa, 100 m, VI.1989, Gauld leg. Paratypes: 1 (INBio): Costa Rica, Heredia Prov., La Selva Biological Station, Finca La Selva, 3 km S Sarapiquí, 50 a 100 m, 1 30.VI.1993, unknown leg., INBio-OET, L_N_268800_535300 #44922; 1 (INBio): same locality, 1 31.VII.1993, unknown leg., Malaise Trap, INBio-OET, L_N_268800_535300 #44715; 1 (INBio): same locality, 1 31.VIII.1993, unknown leg., INBio-OET, L_N_268800_535300 #44717; 1 (INBio): same locality, 1 31.IX.1993, unknown leg., INBio-OET, L_N_268800_535300 #44930. Non-type material (1 ) 1 (CER-UADY): Mexico. Yucatán State, Ría Lagartos Biosphere Reserve, Ría Lagartos Station, Core II, 24.VI 08.VII.2009, A. González-Moreno leg., Malaise trap., dry mounted. Diagnosis This species is very distinctive from all other Mexican species by the following combination of characters: gena less than 0.5 times width of eye in lateral view; fore wing with first abscissa of Cu1a 1.12 1.60 times as long as Cu1 between 1m-cu and cua, and without a pigmented region resembling a vein extending from lower distal corner of first subdiscal cell to hind

50 González-Moreno & Bordera margin of wing; antenna with 27 to 29 flagellomeres; clypeus slightly produced ventrally; eyes strongly convergent ventrally; posterior apex of propodeum extended into a slender, unsculptured neck; fore wing 1m-cu and Cu1a arising from a short common stalk; tergite I of metasoma with spiracle small, not larger than the spiracle of tergite II, and separated from hind margin of tergite by about 1.2 1.5 the dorsal interspiracular distace and scutellum yellow with a median longitudinal black line. This species gives name to the P. tranae species group of Gauld & Bradshaw (1997). HostsR Unknown. DistributionR Neotropical. Brazil, Colombia, Costa Rica, Mexico (Yucatán). HabitatR Gauld & Bradshaw (1997) provided abundant biological data on this species in Costa Rica. In Mexico, P. tranae was found in the savannah-type vegetation. For more detailed description of this habitat see González-Moreno & Bordera (2011). PhenologyR In Costa Rica, this species have been collected abundantly from January to April and sporadically from August to December (Gauld & Bradshaw 1997). In Mexico, it was collected only in June and July on the end of dry season (González-Moreno & Bordera 2011). Podogaster vitticollis (Cresson) (Fig 9) Anomalon vitticolle Cresson 1874: 377. Holotype. Non-type material (4 ) Costa Rica: 1 (AEIC): Guanacaste Province, Santa Rosa National Park, 300 m, XII.1977, Janzen leg.; 1 (AEIC): Puntarenas Prov., Corcovado National Park, Estación Sirena, VII.1977, Janzen leg.; 1 (INBio): Manuel Antonio National Park, 80 m, VII.1991, Zuñiga leg; 1 (INBio), Osa Península, Rancho Quemado, III.1991, IX.1992, X.1993, Saborio, Marín & Gutiérrez leg. Material from Mexico reported by Gauld & Bradshaw (1997) as in the BMNH was not found (G. Broad pers. comm.). Diagnosis Head in dorsal view with gena not swollen, less than 0.5 times width of eye in lateral view; fore wing with first abscissa of Cu1a 0.9 1.1 times as long as Cu1 between 1m-cu and cu-a,1mcu and Cu1a basally separated by a very short abscissa of Cu1, with a pigmented region resembling a vein extending from lower distal corner of first subdiscal cell to hind margin of wing; antenna with 37 to 39 flagellomeres; clypeus slightly produced ventrally; eyes strongly convergent ventrally; mesosoma mainly yellow, with black marks on mesoscutum, propodeum and scarcely on mesopleuron; scutellum coarsely rugose, broadly concave centrally; pronotum laterally and mesopleuron longitudinally striate, with only a smooth area just below the speculum. This species gives name to the P. vitticollis species group of Gauld & Bradshaw (1997). HostsR Unknown. DistributionR Neotropical. Costa Rica, Mexico (Veracruz, Tamaulipas). HabitatR All the Costa Rican specimens were collected on the Pacific Coastal plain and in forests on the Pacific side of the Cordillera mostly at altitudes of less than 500 m (Gauld & Bradshaw 1997). PhenologyR Throughout the year (Gauld & Bradshaw 1997). Acknowledgments We are grateful to Dr. David Wahl and Dr. Hugo Kons (AEIC), Dr. Ronald Zuñiga and Manuel Solis (INBio) and the staff of TMA for permitting us to study Podogaster material during visits to their respective institutions. Dr. Ruíz- Cancino (UAT) borrowed us material of P. eldae and P. ruthae for study. Additionally, Dr. Ronald Zuñiga provided essential data to this work and sent us images of the holotype of P. rosteri. Jason Weintraub (ANSP) sent us photos of the type of P. mexicanus. Dr. Gavin Broad (BMNH) checked paratype material of P. rosteri and corrected the manuscript. Dr. Ilari Sääksjärvi (University of Turku, Finland) took layer photos of P. brunneus, P. lagartensis and P. vitticollis. M. Sc. Luis Salinas Peba took image of P. tranae. Also our thanks to the staff of the Ría Lagartos Biosphere Reserve (Yucatán, Mexico) for allowing access to research facilities and providing authorization for collecting material in this protected area. This study was supported by Projects A/018089/08 and A/026909/09 of AECID (Ministerio de Asuntos Exteriores y de Cooperación, Spain) and by research grants from Consejo Nacional de Ciencia y Tecnología (CONACYT; Mexico) and Conselleria d Educació (Generalitat Valenciana, Spain, Ref: BEST/2010/036). References Cameron P (1886) Hymenoptera, p. 241 328. In Godman FD, Salvin O (eds) Biologia Centrali-Americana or, Contributions to the knowledge of the fauna and flora of Mexico and Central America. Zoology, Volume 1. Cresson ET (1874) Descriptions of Mexican Ichneumonidae. Proc Acad Natural Sci Phila 1873:374 413 Cushman RA (1927) Miscellaneous notes and description of Ichneumon-flies. Proc US Nat Mus 72(2709):1 22 Flores S, Espejel CI (1994) Tipos de Vegetación de la Península de Yucatán. Etnoflora Yucatanense Fascículo 3. Universidad Autónoma de Yucatán, México, p 135 Gauld ID (1976) The classification of the Anomaloninae (Hymenoptera: Ichneumonidae). Bull Nat Hist Mus Entomol 33:1 135 Gauld ID, Bradshaw K (1997) The subfamily Anomaloninae, p.13 176. In Gauld ID (ed) The Ichneumonidae of Costa Rica, 2. Introduction and keys to species of the smaller subfamilies, Anomaloninae, Ctenopelmatinae, Diplazontinae, Lycorininae, Phrudinae, Tryphoninae

Mexican Species of Podogaster 51 (excluding Netelia) and Xoridinae, with an appendices on the Rhyssinae. Mem Am Entomol Ins 57, 485 p. González-Moreno A, Bordera S (2011) New records of Ichneumonidae (Hymenoptera: Ichneumonoidea) from Mexico. Zootaxa 2879:1 21 Graf V (1983) Ichneumofauna do sudeste e sul do Brasil 5 Nova espécie de Philodrymus (Anomaloninae, Hymenoptera). Contrib Am Entomol Inst 20:398 401 Hernandez JV, Osborn F, Herrera B, Liendo-Barandiaran CV, Perozo J, Velásquez D (2009) Parasitoides larva-pupa de Hylesia metabus Cramer (Lepidoptera: Saturniidae) en la región nororiental de Venezuela: un caso de control biológico natural. Neotrop Entomol 38:243 250 Ruíz-Cancino E (2010) Ichneumonidae (Hymenoptera) del Estado de Tamaulipas, México, Serie avispas parasíticas de plagas y otros insectos N. 6. Universidad Autónoma de Tamaulipas, Tamaulipas, México, p 184 Ruíz-Cancino E, Kasparyan DR, Coronado-Blanco JM (2002) 37 Ichneumonidae, p. 631 646. In Llorente Bousquets J, Morrone JJ (eds) Biodiversidad, Taxonomía y Biogeografía de Artrópodos de México: Hacia una síntesis de su conocimiento. CONABIO-ECOSUR-BAYER.