J< u>3 . , J / > ■ r \ /I Journal of Hymenoptera Research Volume 14, Number 2 October 2005 ISSN #1070-9428 CONTENTS AGUIAR, A. P. Cladistic assessment, key and description of two new neotropical genera and species of Gabuniina (Hymenoptera: Ichneumonidae: Cryptinae) 121 BURGE, D. O. Taxonomy, biology, and distribution of seed harvesting ants in the Pheidole californica complex (Hymenoptera: Formicidae) 137 LEATHERS. J. W., D. D. JUDD, AND A. V. Z. BROWER. A review of the species of the new world braconid genus Cyclaulacidea (Hymenoptera) with key and descriptions of nine new species 151 QUICKE, D. L. J. AND N. M. LAURENNE. Notes on host searching by the parasitic wasp Zaglyptogastra Ashmead (Hymenoptera: Braconidae: Braconinae) in Kibale Forest, Uganda 177 QUICKE, D. L. J. AND M. L. STANTON. Trigastrotheca laikipiensis sp. nov. (Hymenoptera: Braconidae): A new species of brood parasitic wasp that attacks foundress queens of three coexisting acacia-ant species in Kenya 182 QUINTERO A., D. AND R. A. CAMBRA T Pappognatha Mickel (Hymenoptera: Mutillidae: Sphaeropthalminae): New species, sex associations, hosts, and new distribution records 191 WHITFIELD, J. B., A. A. VALERIO, AND W. Y CHOI. Redescription of Pelicope yuccamica Mason (Hymenoptera: Braconidae: Microgastrinae), with notes on its unusual biology and relationships 200 NOTE: NIEVES-ALDREY, J. L. AND M. HERNANDEZ NTEVES. OMgtii&mlmanticus Nieves- Aldrey, 1984 (Hymenoptera: Chalcidoidea:,X>rmyridae), a valid\pecies 206 INTERNATIONAL SOCIETY OF HYMENOPTERISTS Organized 1982; Incorporated 1991 OFFICERS FOR 2005 Denis Brothers, President Michael E. Schauff, President-Elect Michael W. Gates, Secretary Justin O. Schmidt, Treasurer Gavin R. Broad, Editor Subject Editors Symphyta and Parasitica Aculeata Biology: Mark Shaw Biology: Sydney Cameron Systematics: Donald Quicke Systematics: Wojciech Pulawski All correspondence concerning Society business should be mailed to the appropriate officer at the following addresses: President, School of Botany and Zoology, University of KwaZulu-Natal, Private Bag X01, Scottsville, South Africa; Secretary, Southwestern Biological Institute, 1961 W. Brichta Dr., Tucson, AZ 85745, USA; Treasurer, PO Box 37012, c/o Smithsonian Institution, MNMH, MRC168, Washington, DC 20013-7012, USA; Editor, Centre for Ecology & Hydrology, Monks Wood, Abbots Ripton, Huntingdon, Peterborough PE28 2LS, UK. Membership. Members shall be persons who have demonstrated interest in the science of ento- mology. Annual dues for members are US$40.00 per year (US$35.00 if paid before 1 February), payable to The International Society of Hymenopterists. Requests for membership should be sent to the Treasurer (address above). Information on membership and other details of the Society may be found on the World Wide Web at http://IRIS.biosci.ohio-state.edu/ish. Journal. The Journal ofHymenoptera Research is published twice a year by the International Society of Hymenopterists, % Department of Entomology, Smithsonian Institution, Washington, D.C. 20560-0168, U.S.A. Members in good standing receive the Journal. Nonmember subscriptions are $60.00 (U.S. currency) per year. The Society does not exchange its publications for those of other societies. Please see inside back cover of this issue for information regarding preparation of manuscripts. Statement of Ownership Title of Publication: Journal of Hymenoptera Research. Frequency of Issue: Twice a year. Location of Office of Publication, Business Office of Publisher and Owner: International Society of Hymenopterists, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560-0168, U.S.A. Editor: Gavin R. Broad, Centre for Ecology & Hydrology, Monks Wood, Abbots Ripton, Huntingdon, Peterborough PE28 2LS, UK. Managing Editor and Known Bondholders or other Security Holders: none. This issue was mailed 7 October 2005 J. HYM. RES. Vol. 14(2), 2005, pp. 121-136 Cladistic Assessment, Key and Description of Two New Neotropical Genera and Species of Gabuniina (Hymenoptera: Ichneumonidae: Cryptinae) Alexandre P. Aguiar Museu de Zoologia, Universidade de Sao Paulo, Av. Nazare 481, Ipiranga, Sao Paulo SP, Brazil 04263-000; email: aguiar.2@osu.edu Abstract. — Phe Cryptini Fenixia n. gen., from the Brazilian Atlantic Forest, and Dineotropica n. gen., from the Amazon basin, are proposed, described, and cladistically compared with literature data for representative species of all genera of the subtribe Gabuniina and 38 outgroup species. A total of 72 species and 51 characters are evaluated. Implied weighting results suggest that Fenixia is closely related to Lagarosoma Gupta, while Dineotropica is closest to Cestrus Pownes. Unweighted analyses were inconclusive, but implied weighting results support the monophyly of Gabuniina essentially as defined in the literature, and suggest that Wuda singularis, and perhaps a few other taxa of Ceratocryptina, might be part of, or a sister taxon of that subtribe. Phe species Fenixia curta n. sp. and Dineotropica lissa n. sp. are described and illustrated. A key to genera of neo- tropical Gabuniina is presented. Phe subtribe Gabuniina was proposed by Pownes (1970) to include a worldwide group of cryptine wasps currently with 31 genera and 309 described species (Yu and Horstmann 1997). Members of this sub- tribe seem to attack xylophagous larvae of Coleoptera and Lepidoptera, exhibiting a highly specialized body structure for this purpose, as noted by Pownes and Pownes (1962): head subspherical; body shape ap- proximately cylindric; ventral tooth of mandible normally longer than dorsal tooth; fore tibiae dilated, having enlarged subgenual organs; antennal tip highly modified; ovipositor compressed, straight and stout, the lower valve with an apical dorsal lobe that encloses most of the tip of upper valve; and subapical metasomal segments enlarged, accommodating spe- cial ovipositor muscles. Phe specialized antennal tip is used to tap the wood, pro- ducing pulses of sound (Pownes and Pownes 1962, Henaut 1990, Often et al. 2000); the echoes are detected with the en- larged subgenual organs (Vilhelmsen et al. 2001, Often et al. 2003), providing infor- mation about the exact location of the host. Broad and Quicke (2000), discussing the adaptive significance of host location by vibrational sounding, further demon- strated that such tibial and antennal spe- cializations are correlated with greater rel- ative host depth, immobility of the host and idiobiosis. Most gabuniines occur in the tropics and subtropics, but there are a few genera confined to the Nearctic region and some others which occur in the Palaearctic re- gion (Gupta and Gupta 1983). Seven gen- era have been described for the Neotrop- ical region, but only two appear to be common, Digonocryptus Viereck, and Agonocryptus Cushman. Phe group has never been extensively studied in South America and the two new genera pro- posed herein point to a much greater di- versity of the subtribe in this region. Comparative studies with Cryptinae, however, are challenging, both because of their extreme diversity, nearing 380 valid genera and over 4500 valid species, and because many of these taxa are also exter- 122 Journal of Hymenoptera Research nally similar, while exhibiting a confusing array of subtle differences, making them difficult to recognize at first sight, compli- cating generic and supergeneric classifi- cation in particular. In an attempt to ob- jectively assess these problems, the pre- sent work uses cladistic analyses with the aim of testing the validity of, and situating the proposed genera into, the tribe Cryp- tini. A final objective is to present a first, while not thorough, cladistic assessment of the monophyletic status of the subtribe Gabuniina based on external morphology, furthering the molecular analyses of Lau- renne et al. (2003) with six gabuniine gen- era, most of them recovered as a mono- phyletic group. MATERIALS AND METHODS General — This work deals exclusively with material acquired through an exten- sive program of field excursions and visits to entomology museums in Brazil, as part of a multi-institutional project developed along the years 2000-2004, now continued by the author in a new program. Speci- mens of neotropical countries unavailable in Brazilian museums were not examined for this work, but are targeted for study as part of the program in progress. Morphological terminology follows Gauld et al. (1997); acronyms for collec- tions follow Arnett et al. (1993). Drawings were prepared by Glaucia Marconato, un- der the author's supervision. Selection oftaxa and characters. — Cladistic analyses were performed exclusively for providing an objective evaluation of the proposed taxa, particularly in relation to published data. Accordingly, character se- lection and coding fit this aim only, and results were not explored for the internal phylogeny of Cryptini or Gabuniina. Literature information was combined with original data, coded into a character matrix, and analysed with cladistic meth- The ingroup includes representatives described Gabuniina genera. A corn- large number of outgroup taxa (38 species) had to be considered, as a re- sponse to the following problems. First, there is a lack of clearly defined sister- groups for Gabuniina and, at the same time, the current subtribal arrangement for Cryptini is highly artificial; with this, outgroup taxa had to be selected from nu- merous species apparently or supposedly related to Gabuniina. Second, extensive tests with different taxa or groups of taxa, during preliminary analyses, showed that reasonably stable results could only be ob- tained with a large number of outgroup taxa. The character set has a slight emphasis on features habitually used for describing genera of Gabuniina. Many characters were coded directly from illustrations, and checked with the corresponding descrip- tions whenever possible. The considered taxa are described and illustrated in Townes (1970), Gupta and Jonathan (1970), and Gupta and Gupta (1983). Char- acters 1-2, 4, 6-8, 18, 30, and 40-43 (Table 1) were coded from the general descrip- tions of Townes (1970) for the genera. For these characters, all species of a given ge- nus were coded with the same character state. Although this may contribute, dur- ing tree search, to species in the same nominal genus to end up grouped in one clade, therefore supporting the original concept, this scheme was ultimately adopted because the potential problem is only marginally relevant for the purpose of this study. Character 7, the percentage of variation of the fore wing length, was calculated by taking the difference be- tween the largest and the smallest wing length registered for the genus, and divid- ing it by the smallest wing length value. Two apparently distinct tendencies (Fig. 1) were interpreted and coded as two dis- tinct character states. Regression lines for Fig. 1 were calculated with smoothing splines (Venables and Ripley 1997), which draw the curve that best suits a given data set. Phylogenetic analysis. — Tree searching Volume 14, Number 2, 2005 123 was performed with heuristic analyses in NONA, version 2.0 (Goloboff 1993b) aided by Ratchet (Nixon 1999a), and with im- plied weighting in PIWE, version 2.8 (Go- loboff 1997), which resolves character con- flict in favor of characters that have less homoplasy during tree search. Cladogram analysis was performed with WinClada, version 1.00.08 (Nixon 1999b), which also incorporates the program Ratchet. All multistate characters were first treated as unordered, then characters 20, 32 and 33 were reinterpreted from the initial trees, and run as ordered. At this stage, changes for character 20 were interpreted as 0«— 1^>2 and the character was respective- ly recoded in the matrix as l«-0— >2, to al- low the respective changes to be accurate- ly considered during tree searching. For Ratchet, independent searches were performed with a sample of 5-8 charac- ters, and 3000 iterations on each run. The resulting trees were submitted at once to NONA, screened with best and unique, which discard non-optimal trees and trees that are optimization-sensitive, and then submitted to increasingly exhaustive searches, as follows: swapping with max*, a procedure which also certifies that the trees found will belong to a "complete is- land" (cited from PIWE manual), and fur- ther swapping with ms*l, and jump*l to jump*4, which search for better trees in different "islands" by generating slightly less optimal trees from the ones found be- fore. All steps found more or better trees, except jump*4. For PIWE the options "hold 10000, mult*100" were used, with resulting trees submitted to further swapping exactly as described above for NONA. Searches were performed with the default value for K, the constant of concavity (Goloboff 1993a) and for K = = 2 and K ' 1. Higher values of K (4-6) were not tested because the intention was to check the maximum influence that less homoplasious charac- ters could have on the phylogeny. RESULTS Table 1 shows the character list and character state coding, and Table 2 pre- sents the respective character matrix. Searches with NONA /Ratchet found 6531 most parsimonious trees of 560 steps, Ci 14, Ri 54. The respective strict consensus tree is almost entirely collapsed and only marginally informative for Gabuniina. For this reason, it will not be considered here. With implied weighting, results are as fol- lows: searches with jK = 1 found 3071 trees of fit 110.4; K = 2 generated 3 trees of fit 141.9; and K - 3, the default value, yield- ed 29 trees of fit 172.9. Results with Kl are not illustrated because they were gener- ally similar to those obtained with K2-3, except that the correspondent consensus tree was considerably less well resolved, with 34 collapsed nodes versus 1 and 11 collapsed nodes for K2 and K3 trees, re- spectively. All cladograms for each of these searches preserved clades of interest for Gabuniina as a whole (Figs. 2, 5), and for the proposed new genera (Figs. 3-4, 6- 7). All weighted searches recovered a clade with 30 Gabuniina genera, supported by a single, non-homoplasious character state (42:1, petiole spiracle approximately at middle). Relationships among the taxa within Gabuniina were also similar in these searches. Fenixia n. gen. appears in a collapsed clade on searches with XI, but is associated with Lagarosoma on all searches with K2-3 (Figs. 3, 6); Dineotw- pica n. gen., in its turn, was recovered forming a clade with Cestrus on all weighted searches, supported by at least two synapomorphies (33:0, hind wing vein M+Cu weakly convex, and 51:1, ovi- positor subapically with a microsculptu- red area) (Figs. 4, 7). Characters traditionally used to define Gabuniina (numbers 4, 42, 48 in Table 1) performed similarly in each analysis (Figs. 2, 5, arrows), generally showing compar- atively high Ci and Ri values for the re- 124 Journal of Hymenoptera Research Table 1. Character coding. Abbreviation: n/a, non-applicable. jsj0 Description 01 Clypeal margin, number of median teeth 0 [0], 1 [1] or 2 [2], even if small 02 Mandible with ventral tooth longer than dorsal [0]; equal size [1]; ventral tooth shorter than dorsal [2] 03 Epomia short and weak [0]; long and strong [1]; represented by a group of wrinkles, rather than a single carina [2]; absent [3] 04 Fore tibia of female regular-looking [0]; swollen, basally constricted [1] 05 Sternaulus complete, reaching middle coxa [0]; incomplete, reaching 0.45-0.65 of the distance to mid- dle coxa [1]; absent [2] 06 Pleural carina absent [0]; distinct and complete [1]; distinct but weak and incomplete [2] 07 Fore wing length variation up to 125% [0]; more than 125% [1] 08 Fore wing vein 1-Rs + M straight or slightly convex [0]; concave or sinuous, even if slightly [1] 09 Fore wing crossvein lm-cu about as long as vein 1-Rs+M [0]; distinctly shorter [1]; distinctly longer [2]; n/a (limit between veins indistinct) [-] 10 Fore wing bulla on vein 1-Rs + M central [0]; apical, reaching cell l+2Rs or nearly so [1]; bulla absent [2] 11 Fore wing crossvein lm-cu straight or uniformly curved [0]; sinuous or somewhat irregular [1] 12 Fore wing without a short vein projection (ramellus) arising at meeting of veins lm-cu and Rs + M [0]; a short projection present [1] (partially linked to character 14) 13 Fore wing limit between lm-cu and 1-Rs + M distinct [0]; indistinct, veins perfectly continuous [1] 14 Fore wing vein lM+Rs weakly and uniformly curved, or straight [0]; slightly sinuous ot weakly ir- regular [1] 15 Fore wing crossvein lcu-a usually far from base of lM + Rs (basad by more than 0.1 its own length) [0]; veins very close (approximately basad by 0.1 or apicad) or opposite [1] 16 Fore wing crossvein lcu-a at approximately 90° with M + Cu [0]; distinct obtuse angle [1]; distinct acute angle [2] 17 Fore wing vein 2-Cu distinctly longer than crossvein 2cu-a [0]; nearly of the same length or 2cu-a slightly longer [1]; 2cu-a much longer than 2-Cu [2]; 2-Cu entirely absent [3] 18 Fore wing vein 2-Cu and 2cu-a aligned [0]; angled, even if slightly [1]; n/a [-] 19 Fore wing vein 4-Rs uniformly curved [0]; sinuous or irregular [1] 20 Fore wing crossvein 2m-cu with bulla mostly central to mostly ventral [0]; placed entirely or mostly on anterior 0.5 [1]; nearly reaching or reaching cell l+2Rs [2]. Recoded in the matrix as 1, 0, 2 respectively, and run as additive. 21 Fore wing cell l+2Rs (areolet) size, even if open, small [0]; large, about as tall as width of pterostig- ma [1]; not differentiated [-] Fore wing crossveins 2r-m and 3r-m parallel or nearly so [0]; distinctly but slightly or moderately convergent towards anterior margin of wing [1]; strongly convergent [2]; n/a (areolet open or not differentiated) [-] 23 Fore wing veins 2-M and 3-M approximately of the same length, or one slightly shorter than the other [0]; 3-M distinctly longer than 2-M [1]; 2-M distinctly longer than 3-M [2]; n/a [-] 24 Fore wing crossvein 3r-m tubular, normal [0]; entirely or partly nebulous or spectral, including "with bulla" [1]; not differentiated, cell l + 2Rs open [2]; n/a (cell l+2Rs not developed) [-] 25 Fore wing cell l+2Rs pentagonal, or nearly square or circular, even if slightly taller than wide or if open [0]; transversely elongate [1] 26 Fore wing vein 4-M slightly to distinctly longer than 4-Rs [0]; nearly as long as, or shorter, than 4-Rs [ 1 1; 4-M nebulous or spectral on apical half or more [2] Hind wing vein 2-1 A short, not reaching wing margin, or absent [0]; ending near or at wing margin [1] Hind wing vein 1-Cu with nearly the same length as crossvein cu-a [0]; 1-Cu distinctly longer [1]; 1-Cu distinctly shorter [2] Hind wing vein 2-Rs entirely tubular [0]; apical half or more nebulous or spectral [1] 1 find wing crossvein lr-m entirely tubular [0]; with one bulla 1 1 ] wing veil s 1-Rs and 2-Rs forming a distinct angle (cell Rl somewhat trapezoidal basally) [0]; continuous or nearly so (cell Rl pointed or lanceolate basally) [1] [coded as multi-state when lubtful or intermediate] Cu and M at about 90° [0]; forming a distinctly acute angle [1]; forming a distinct- obtuse angle [2] Volume 14, Number 2, 2005 125 Table 1. Continued. Description 33 Hind wing vein M+Cu uniformly and weakly convex, or straight [0]; strongly convex [1]; concave [2] 34 Hind wing vein 1-R1 (the short section of Rl detached from wing margin) distinct [0]; not differenti- ated [1] 35 Anterior transverse carina of propodeum straight or weakly and uniformly curved [0]; strongly curved, trapezoidal or acuminate medially [1]; fused with posterior transverse carina [2]; absent [3] 36 Posterior transverse carina of propodeum present, even if interrupted centrally or indicated only by lateral crest or spines [0]; entirely absent [1] 37 Posterior transverse carina of propodeum uniformly convex, weakly or strongly, even if briefly inter- rupted centrally [0]; strongly bell-shaped or trapezoidal [1]; forming lateral crests [2]; forming later- al tubercles or spines [3]; n/a (absent) [-]. 38 First metasomal tergite short and triangular, length/(maximum width - miniumum width) less than 4.0 [0]; regular, somewhat elongate lg/(wmax - wmin) 4.0-6.0 [1]; long and slender, lg/(wmav - wmin) over 6.0 [2] 39 First metasomal tergite without a basolateral triangular tooth [0]; tooth present, even if vestigial [1] 40 First metasomal tergite without an extra basolateral triangular tooth [0]; extra tooth present [1] 41 First metasomal tergite without dorsolateral carina [0]; partially developed [1]; complete [2] 42 Spiracle of first metasomal tergite placed beyond middle [0]; at or basad of middle [1] 43 T7-8 in lateral view of similar size or shorter than T5-6 [0]; distinctly wider [ 1 ] 44 Upper valve of ovipositor in lateral view distinctly widest preapically, the nodus tall, giving triangu- lar aspect to apex [0]; width decreasing uniformly, nodus weak or not evident, apex not triangular [1] 45 Upper valve of ovipositor apically straight or nearly so [0]; distinctly downcurved [1]; distinctly up- curved [2] 46 Upper valve of ovipositor apically without serrations [0]; serrations present [1] 47 Upper valve with preapicaJ notch [0]; absent [1]; modified structure [2] 48 Lower valve of ovipositor apically regular, not dilated [0]; dilated and overlapping upper valve as a lobe [1]; apically widened to cover entire tip as a sheath [2] 49 Lower valve of ovipositor with serrations along entire tip [0]; restricted to the very tip, or serrations absent [1] 50 Ovipositor tip with upper valve apex blunt or only moderately pointed [0]; ending in a long and narrow point [1] 51 Ovipositor just basad of apical teeth smooth and polished [0]; with a distinctly microsculptured area [1] spective trees, as follows. Character 4 (swollen fore tibia of female) with Ci 12- 14 and Ri 78-81; character 42 (Tl with spi- racle at or behind middle) with Ci and Ri = 100 in all implied weighting clado- grams; and character 48 (lower valve with lobe enclosing upper valve) with Ci 33 and Ri 87 also in all cladograms. DISCUSSION Preferred cladograms. — The ambiguous results obtained with K\ searches point to a negative consequence of the maximized weight given to a few, less homoplasious characters in the matrix. This clearly de- preciated most of the already weakly in- formative characters, to a point where K\ searches could have mimicked unweight- ed searches, which were mostly uninfor- mative. Cladograms obtained with K2-3 seem therefore to represent the best pos- sible results with the available information in the character set of Table 2, and because of this are adopted as the preferred phy- logenetic interpretation. Gabuniina. — In spite of a few differences at the base of the respective clade, the weighted analyses generally corroborate the idea that the Gabuniina of authors might be a monophyletic group. They also 126 Journal of Hymenoptera Research 20 30 Genera Fig. 1. Plotted data for character 7 (Table 1), with regression lines showing two distinct tendencies (concave vs. convex lines) for the known percentage of variation of the fore wing length, for the genera listed in Table 2. generally confirmed the relevance of char- acters traditionally used in the literature (numbers 4, 42, and 48, Table 1) to define and characterize the subtribe, lending sup- port to Gabuniina plus a few basal taxa (particularly Ceratocryptina), or to Gabu- niina minus a few of its basal taxa. The marked phylogenetic importance of these characters is also suggested by their com- paratively high values of Ci and Ri. The repeated presence of W. singularis, and other Ceratocryptina as well, at the base of the Gabuniina clade suggests that this group, or possibly the genus Wuda as a whole, may represent a basal member of Gabuniina, or its sister taxa. The implied weighting results also support the sugges- tion of Gupta and Gupta (1983) that the genera of Gabuniina "appear to form two distinct groups" based on the presence/ absence of the pleural carina (ch. 6, Table 1). This character showed high values of Ci (66-100) and Ri (91-100) on all clado- grams, suggesting that Gabuniina with a pleural carina may represent a monophy- j;roup. Gupta and Gupta (1983) also milar groups based on the number of clypeal teeth (ch. 2), and on the comparative length of hind wing vein 1-Cu and crossvein cu-a (ch. 28), but this is not supported by the current analyses, with very low Ci and Ri values for both these characters. Relationships of Fenixia and Dineotropi- ca. — Results clearly indicate that these taxa must be assigned to Gabuniina, as de- fined in the literature and as recovered here. Fenixia n. gen. and Dineotropica n. gen. also appear to be only distantly re- lated to one another, each being recovered within its own large and distinct clade, and therefore isolated from each other by numerous steps. At the same time, Fenixia seems most closely related to Lagarosoma Gupta, the two forming a clade supported by an identical set of synapomorphies (Figs. 3, 6). Such relationship is further corroborated by the fact that both these genera are known exclusively from south- eastern Brazil. The close relationship of the central Amazonian Dineotropica with the essen- tially Mexican Cestrus seems evident by the sharing of a unique feature within Ga- buniina, the ovipositor apex with a mi- crosculptured area just basad of apical teeth (ch. 51:1), as well as the hind wing with a weakly convex vein M+Cu (ch. 33: 0), and the propodeum with posterior transverse carina strong, complete. None- theless, the uniqueness of Dineotropica is evident by at least 9 apomorphies, repeat- edly recovered for this genus in all Kl-3 cladograms (Figs. 4, 7). DESCRIPTIONS OF NEW TAXA Fenixia Aguiar, n. gen. Figs. 8-12 Type species. — Fenixia enrta Aguiar, by monotypy and present designation. Description. — Fore wing 6.3-10.7 mm long. Frons finely granulose, with a short median carina developed near anterior ocellus only. Clypeus weakly convex, more projected ventrally, the apex trun- Volume 14, Number 2, 2005 127 cate and with a median tooth. Mandible epomia small (vs. long and strong), fore 1.5 as long as basal width, dorsal tooth as wing vein 4-Rs sinuous (vs. straight), cell long as ventral tooth. Occipital carina l+2Rs large and pentagonal, 2r-m and meeting hypostomal carina. Epomia short 3r-m convergent, 2-1 A not reaching wing and weak. Sternaulus sharp and reaching margin, and ovipositor sheath very short, middle coxa. Epicnemial carina distinct 0.7 as long as hind tibia (vs. 1.1 as long), along entire height of mesopleurum. Hind According to the cladistic analyses, Fenixia margin of metanotum without projections also seems related to the sympatric genus on each side of postscutellum. Area be- Lagarosoma Gupta, from which it can be tween metanotum and propodeum mod- isolated by the clypeus margin with one erately deep, narrow, forming a polished tooth (absent in Lagarosoma), fore wing smooth trough. Propodeum as long as 2-Cu distinctly longer than 2cu-a (vs. wide. Propodeal spiracle oval elongate, much shorter), crossvein lcu-a very close Pleural carina absent. Propodeum in front or opposite lM + Rs (vs. far from base), cell of basal carina punctate and allutaceous; l+2Rs higher than wide (vs. wider than behind basal carina transversely rugulose high), 2r-s and 3r-s about same length (vs. to rugose. Juxtacoxal carina absent. Apical 3r-s distinctly longer), and basal carina of carina of propodeum indistinct, or indi- propodeum strong and distinct (vs. indis- cated by weak lateral crests. tinct or absent). Other distinctive character Fore wing cell l + 2Rs about as long as states are indicated on the respective clad- width of pterostigma, a little higher than ogram (Figs. 2, 5) and key to neotropical wide, cross veins 2r-m and 3r-m distinctly genera, below, convergent, about same length. Ramellus absent. Crossvein lcu-a slightly but dis- Fenixia curta Aguiar, n. sp. tinctly basad of vein lM + Rs. Hind wing JH18S- o-iz vein 1-Cu 1.9 length of crossvein cu-a; 2- Description. — Female (holotype). Fore 1A reaching 0.5-0.8 the distance to wing wing 10.7 mm long. Clypeus weakly con- margin. Fore tibia in female moderately vex, more projected ventrally, the apex swollen. Fourth segment of all tarsi deeply truncate and with a median tooth. Man- bilobed. First metasomal tegite with a dis- dible teeth of equal length. Occipital cari- tinct lateral triangular tooth at the base; na low and sharp throughout, joining the dorsolateral carina absent; spiracle exactly weakly raised hypostomal carina below, at middle; sternite about 0.43 the length of Pronotum: epomia weak, sharp, and short, tergite. Ovipositor 1.2 as long as hind tib- distinct only in between dorsal and ven- ia, projecting beyond metasoma for half of tral yellow marks; area behind epomia, in its own length; ovipositor sheath about between yellow stripes, with longitudinal 0.65 as long as hind tibia. Lower valve of rugulosity. Mesonotum: notauli deep, con- ovipositor with a weak to distinct subapi- verging posteriorly, blending with longi- cal lobe that partly encloses upper valve; tudinal rugulosities on and behind central apex with 8 teeth. yellow spot; notauli and rugulosities end- Etymology. — A reference to the city of ing far from scuto-scutellar groove. Scu- Fenix (Parana, Brazil), the collecting local- tellum micropunctate. Mesepisternum ity for one of the paratypes. finely obliquely strigate, stronger dorsally; Comments. — The genus runs to Dagathia epicnemial carina entirely distinct; ster- Cameron in the key provided by Townes naulus strongly sinuous, distinct from (1970) for the world genera of Gabuniina, epicnemial carina to base of hind coxa; but can be isolated from this Oriental ge- without any indication of a depression be- nus by having mandible teeth of equal tween sternaulus and speculum. Meta- size (vs. ventral tooth a little longer), pleuron densely rugulose. Propodeum 128 Journal of Hymenoptera Research Table 2. Character matrix for selected taxa of Cryptini. Subtribes (Subt): Agrt, Agrothereutina; Bare, Bar- yceratina; Cert, Ceratocryptina; Coes, Coesulina; Cryp, Cryptina; Gabn, Gabuniina; Glod, Glodianina; Gory, Goryphina; hymn, Lymeonina; Meln, Melanocryptina; Mest, Mesostenina; Ospr, Osprynchotina. Polymorphism: a, 01; b, 12;c,02;d, 03. Subt. Species 1 0 Gabn Agonocryptus discoidaloides 1071001100 1001001000 1021001001 lOlaOOlOOO 0111001100 0 Gabn Ahilya bicornigera 1101010??? ?????????? 00?00??0?? ????0a??10 0111101100 0 Gabn Amrapalia multimaculata 10010000-? 0010001100 1100011100 020??01010 bl???????? 0 Gabn Anepomias splendidus 10011070-1 0011011000 1100001000 lla001-100 0011001100 0 Gabn Apocryptus praeciarus 2111110000 0000111000 1001001101 0000100100 2110001100 0 Gabn Arhytis maculiscutis 1011000100 0011001001 lOOlOllcOl 12a001-010 alllOOOlOO 0 Gabn Cestrus calidus 1111011010 0000010001 1201001101 0000100010 0111000100 1 Gabn Cryptohelcostizus alamedensis 0011200070 1011001010 1201011001 a00a01-000 0111001100 0 Gabn Dagathia multimaculata lllla001-0 0000001011 1020011100 1100002010 alllOOHOO 0 Gabn Digonocryptus crassipes badl0110-0 0011101101 1001001100 1010100010 alllOOHOO 0 Gabn Dineotropica lissa n. sp. 00010171-0 0011101010 1021011001 1101000100 0111007100 1 Gabn Dinocryptus niger 10a0a20112 1001001000 1021011000 0000200010 1111001100 0 Gabn Eurycryptus fondamentalis 00110000-2 0010001010 1001711000 1010000010 0011011100 0 Gabn Fenixia curta n. sp. 11010000-0 0011011011 1101000101 10110a2010 0111000100 0 Gabn Gabunia ruficoxis 1011200020 0001011100 1211011000 a00001-100 0111001100 0 Gabn Gerdius cinctus OldOOlOlOO 0000021101 0017011001 lOaOOOOOlO 2111000100 0 Gabn Hackerocryptus dentatus 1137707110 0000011100 0—2-021?? ?????l-000 7171001000 0 Gabn Hadrocryptus sp. al01007000 1001001001 1001011100 1100102010 0111000100 0 Gabn Kriegeria heptazonata 1111010110 0001102110 OOOOOOlcOl 121011-010 clllOOOlOO 0 Gabn Lagarosoma assitum 023100001? 0007102110 112110017? ?71?01-100 0111007100 0 Gabn Lophoglutus bouceki 1037000100 0000111110 0 — 200117? ????102010 7071001100 0 Gabn Microstenus canaliculars 0011000000 0010001100 0--2011000 0010000010 2011011100 0 Gabn Nesolinoceras ornatipennis 10010070-1 0010010002 1201001001 001001-000 0111001100 0 Gabn Pharzites sp. 0121000000 1000010000 1111011200 001001-010 0111001100 0 Gabn Prosthoporus terani 020111???? ????0???7? 101117227? ?71?01-?00 0111001100 0 Gabn Pterocryptus uchidai 10110100-0 0011001100 1101011200 a010002000 0111001100 0 Gabn Schreineria annulata OOdllOOlOO 0001002000 0002011200 a20001-011 0111007100 0 Gabn Spathacantha apicallis 1131017110 0001002101 1120001200 00a001-011 2111000100 0 Gabn Tanepomidos assamensis 121101001? 000010111a 0001001000 021?00aal0 2101000100 0 Gabn Torbda geniculata bOllbOlOOO 1017012000 1121111000 10a011-000 1111001100 0 Gabn Trypha atriceps 01310000-0 0010001000 1001001001 101031-100 OlOlOOlaOO 0 Gabn Xanthocryptus vesiculosus lOdlaOHOO 0001011001 0020001000 011001-010 0111001100 0 Gabn Xoridesopus sp. 1000011100 0001001000 1001011001 001001-010 2111001100 0 Gabn Xoridesopus verticalis 1001011100 0001001000 1117011000 0010000010 2111001100 0 Agrt Agrothereutes abbreviatus* 01a01?l- — 1— —102010 1000000000 0 Agrt Agrothereutes sp. 01a01?1001 1100100002 1001000101 0011102010 1000000000 0 Agrt Gambrus incubitor 11301710-1 0010100002 1001021100 1011000010 1000000000 0 Agrt Trychosis neglecta 1110171000 1000101100 1000001001 0001001007 7000000000 0 Bare Baryceros texanus 7210a?1100 1000101011 0-22011100 a011103000 0011111000 0 Cert Aprix nutatorius 0130077100 0001100111 0020001101 ala0002110 7000001000 0 Cert Ceratocryptus bituberculatus 00300701-0 1011000111 0020101100 0001000200 0000000001 0 Cert Chamula reliqua 0110070010 0011001010 0000001100 lOaalOOHO 0007?????? 0 Cert Lorio austerus 11101clll2 0000001110 0000011101 al00002110 7010001000 0 Cert Trachyglutus polychromus 1101077110 0000101111 0-02002101 1011102000 1011001000 0 Cert Wuda singularis 0130170110 0000111010 0 12101 0110000010 0011001100 0 Coes Coesula fulvipes 01a0070000 0000101110 0-02001101 OOlOlOaOlO 2000000000 0 Cryp Caenocryptus shikokuensis 0170071000 1100101011 1101001101 lOaOlOOOOO 2000000001 0 Cryp Dotocryptns bellicosus 0070070021 1001100010 1000001000 0001101100 0007001200 0 Ischnus inquisitorius ?1?01?1010 1000101112 1201000100 a011002010 1000000001 0 Lanugo retentor 0170171000 1000110011 1001001101 0001001010 2000000001 0 Volume 14, Number 2, 2005 129 Table 2. Continued. Subt. Species Cryp Cryp Cryp Cryp Cryp Glod Gory Gory Gory Gory Gory Gory Gory Gory Lymn Lymn Lymn Lymn Meln Meln Mest Ospr Trachysphyrus cleonis Trachysphyrus cyanipennis Trachysphyrus lachnaels Trachysphyrus tucman Xylophrurus dispar Glodianus sp. Buodias longidentatus Buodias ruficoxis Biodias sp. Goryphus basilaris Goryphus communis Listrognathus pubescens Necolio imperialis Necolio sp. Bicryptella vera Lymeon orb us Pachysomoides fulvus Polycyrtidea flavopicta Melanocryptus sp. A Melanocryptus sp. B Mesostenus transfuga Osprynchotus gigas 01?00?1021 1001100012 1201001a01 a020103000 1000000000 0 01?00?1022 1100110012 lOOOOOlaOl al00103000 1001101000 0 01?00?10-0 0011100012 1201011a01 0001103000 1000000000 0 01?00?1001 1100100012 lOHOOOaOl aOOOlOOOOO 1000200001 0 10?10?0000 0100100111 1201001001 OOalOOOOOO 2001000100 0 0?100?0120 0001101011 0020001100 0011001200 0001011001 0 ?la00?0021 0000101112 1001001101 00a0003110 2000000000 0 ?la00?1100 0000111111 1021111101 a0a0102110 2001001000 0 ?la00?1020 1070101111 1021001100 12a001-110 2000000011 0 all00?1000 0000100111 1001071101 a010002010 2000000000 0 all00?1100 0000101111 0001001101 a01a002010 2000000000 0 ?1101?1120 0000000011 1021001101 OOlalOlOlO 2001002000 0 01d01?1100 0001110111 1021001101 a0a0102110 aOOOOOOOll 0 OldOlPllOO 0001100111 1020001100 lOalOOlllO aOOlOOOOlO 0 0bl00?00-0 0010100100 0000001101 a21a!00000 7001101000 0 2b?10?1100 1000102100 1-02000101 1211002000 0000000000 0 0bl00?0100 0000110010 0-02001101 0210101000 2000000011 0 0bl01?0010 0001103-10 0-02002-11 1011000200 0000000001 0 1200171100 0000100110 1101001101 0000103100 lOlaOOlOOO 0 1200171100 0001111111 1111001101 a010002100 1010001000 0 0110071100 OOlOaOOOll 0-221clll0 0000100110 7001000010 0 0011070110 0101121011 1211011200 000011-270 7001000200 0 * Corrected spelling for Agrothereutes abbrcviator according to Horstmann (2001). scarcely pilose; anteriorly shallowly punc- tate near anterior transverse carina, the punctures becoming progressively more scarce towards axillary trough; this area also distinctly and densely alutaceous; area posterior to transverse carina trans- versely rugose; spiracle weakly oval, al- most circular; anterior transverse carina low, straight, except weakly arched cen- trally; propodeal apodeme represented by low carina on center of lateral yellow spot; pleural carina indistinct. Legs: all preapi- cal tarsomeres deeply bilobed. Metasoma: Tl dorso-lateral carina distinct only from spiracle to apical margin; Tl-4 very finely microsculptured, much stronger and matt on T2; then gradually changing from alu- taceous on T5 to almost polished smooth on T8; T8 weakly to strongly convexly folded centro-longitudinally. Ovipositor sheath dilated and spoon-shaped on its apical 0.3; lower valve of ovipositor with 8 apical teeth; upper valve with tiny sub- apical notch. Wing venation as described for the genus. Color. — Head and mesosoma black with yellow marks; metasoma reddish. Head: scape, pedicel, flagellomeres 1-3, and bas- al 0.8 of 4th, black; apex of 4th, 5-8 entirely, and most of 9th, yellow; 10-13 dark brown, 14 to apical entirely brown; 28 flagello- meres total on both antennae, the apical one moderately compressed and protrud- ed at apex. Orbital band yellow, complete except very briefly interrupted at bottom of eye; taking entire width of gena ven- trally, gradually narrowing dorsally to as narrow as 0.4 the distance between eye margin and occipital carina at temple; width at frons and face about 0.25 intero- cular distance; face mostly yellow, partly fusing with yellow of orbital band; clype- us yellow, with wide black area isolating it from yellow of face and orbital band; also yellow on labrum, large spot taking basal 0.4 of mandible, and labial and max- 130 Journal of Hymenoptera Research GLOD Glodianus sp. BARC Baryceros texanus 21243] GABN Eurycryptus fondamentalis GABN Microstenus canaliculars GABN Lophoglutus bouceki CERT Wuda singularis 8 10 13 1421 2251 38 GABN Fenixia curta n. sp. GABN Anepomias splendidus GABN (30 Genera) 0 2 1 10 3 8 1*. 17 20 22 2'. 28 35 GABN Lagarosoma assitum GABN Cestrus calidus GABN Dineotropica lissa n. sp. 7 21 24 39 4 DOOOD — 0 0 0 0 0 - GLOD Glodianus sp. BARC Baryceros texanus - GABN Eurycryptus fondamentalis -GABN Microstenus canaliculars CERT Chamula reliqua CRYP Dotocryptus bellicosus CERT Trachyglutus polychromus GABN Lophoglutus bouceki 26 27 0 I I 1922 2627 33 I I 0 I GABN Fenixia curta n. sp. 17 18 20 23 25 38 39 GABN Lagarosoma assitum o i i i : GABN Cestrus calidus GABN Dineotropica lissa n. sp. CERT Wuda singularis - H 1013 142122 3138 GABN Anepomias splendidus GABN (30 Genera) Figs. 2-7. Summary of clades containing the Gabuniina of authors, and clades containing the genera Fenixia and Dineotropica, preserved on trees obtained with implied weighting searches. 2, Clades from the strict consensus of 3 trees obtained with searches for K = 2. 3, Same, for section of tree containing Fenixia. 4, Same, for Dineotropica. 5, Clades from the strict consensus of 29 trees obtained with searches for K = 3 (default value). 6, Same, for section of tree containing Fenixia. 7, Same, for Dineotropica. The arrows indicate characters traditionally used to define the subtribe Gabuniina. Subtribe abbreviations: BARC, Baryceratina; CERT, Cer- atocryptina; GABN, Gabuniina; GLOD, Glodianina; LYMN, Lymeonina. illary palpi, except for brown apical arti- cle. Mesosoma black, except yellow as fol- low: broad band on pronotum along an- terior and ventral margins, ending dis- tinctly before reaching posterior corner, barely interrupted centro-anteriorly, and a moderately wide band on central 0.7 of dorso-lateral margin; diffused marks on prosternum basally; large hexagonal spot taking about 30% of mesopleuron; all sub- alarum and tegula; small spot dorsally on 'culum; central subcircular spot on me- in between area of convergence cutellum and post-scutellum entirely; axillary trough except small area mesally; large subtriangular spot on each side of propodeum, behind posterior transverse carina, each one about 0.3 as wide as propodeal width. Legs: fore and mid coxae, except mesally, pale yellow, large spot dorso-basally, at level of dorsal articulation of hind coxa, yellow; hind coxa, trochanter and femur red-brown; fore and mid trochanters, femora, and bas- itarsomere yellowish brown, with blackish centrally; all tibiae and tarsomeres 1-4 of hind leg golden yellow; tarsomeres 3-5 of fore and mid legs, and apical tarsomere of Volume 14, Number 2, 2005 131 ^s» 15 16 Figs. 8-18. Fenixia carta n. sp. Holotype 9: 8, Habitus, left. 9, Propodeum, dorsal. 10, Tip of ovipositor, left. Paratype 9: 11, Right fore wing. 12, Right hind wing, same paratype. Figs. 13-18. Dineotropica lissa n. sp. Holotype 9. 13, Habitus, left. 14, Propodeum, dorsal. 15, Ovipositor and sheath, left. 16, Tip of ovipositor, left. 17, Right fore wing. 18, Right hind wing. Drawings by Glaucia Marconato. 132 Journal of Hymenoptera Research hind leg, dark brown. Metasoma dark red, basal segments darker than apical seg- ments; corners of Tl yellowish. Ovipositor dark red; sheaths dark brown. Wings hy- aline. Male. — General morphology and color similar to female, except for yellow tones clear and bright; orbital band not inter- rupted; face and clypeus entirely yellow; speculum entirely yellow; Tl apical mar- gin with distinct yellow stripe. Male from Agua Funda with yellow marks also on mesosternum, near base of coxa and along sternaulus. Material— 15 9 9,236. Holotype 9 DZUP "N. Teutonia-S.C, Brasil-XI/1967, F. Plaumann leg." Paratypes: DZUP: 9 9, "N. Teutonia, SC, Brasil-XI/ 1967, F.Plaumann leg."; "P. Grossa (Vila Velha), PR, Reserva IAPAR, BR 376, Brasil, 15.IX.1986, Lev. Ent. PROFAUPAR, Malaise" (3 specimens); same, 06.X.1986; same, 11.VIII.1986; "Jundiai do Sul, PR, Fa- zenda Monte Verde, Brasil, 24.XI.1986, Lev. Ent. PRO- FAUPAR, Malaise" (2 specimens); same, 15.XII.1986; "Sao Jose dos Pinhais, PR, Serra do Mar, BR 277, Km 54, Brasil, 16.111.1987, Lev. Ent. PROFAUPAR, Mal- aise"; "Curitiba, PR, CI, 28.XII.1976-10.I.1977, V. Graf leg."; "Curitiba, PR, Brasil, 09. XI. 1978, F. Giacomel leg"; "Colombo, PR, EMBRAPA, BR 475, Km 20, Brasil, 22.IX.1986, Lev. Ent. PROFAUPAR, Malaise"; 6, "Fenix, PR, 03.X.1986, A. F. Kumagai col"; 9, "Blumenau, SC, Brasil, V. Graf leg.". MZUP: 6, "Sao Paulo, Agua Funda, SP, 21.VIII.1967, Curso D.Z." Variability. — General morphology: ru- gulosity laterally on pronotum sometimes very weak; propodeal apodeme some- times entirely absent, not even indicated as a low carina; metapleuron sometimes densely pilose. Color: Yellow may change to pale yellow or almost white in small specimens; extension of yellow on face and clypeus highly variable, from entirely and continuously yellow in female from Monte Verde (15.XII) to widely black in between these areas, forming an "M" or "H" figure, as in female from Blumenau. Presternum entirely black to mostly yel- low; yellow on speculum varying from small spot dorsally to yellow on its 0.8 ipe and size of yellow area on on highly variable, taking up to 1 area. On smallest female examined (from Monte Verde, 24. XI), black replaced by dark brown, and hind coxa without yellow spot. Female from Sao Jose dos Pinhais with ovipositor sheath yellow on inner surface. Etymology. — From the Latin curtus, meaning short; in reference to the length of the ovipositor sheath. Distribution records. — Southern and southeastern Brazil (SC, PR, SP). Dineotropica Aguiar, n. gen. Figs. 13-18 Type species. — Dineotropica Jissa Aguiar, by monotypy and present designation. Description. — Fore wing 16.1 mm long. Frons smooth, with a short median carina developed centrally only. Face with a strong U-shaped fold from one antennal foramen to the other. Clypeus flat, apical 0.3 truncate, apical margin uniformly blade-like, translucent, without a median tooth. Mandible 1.67 as long as basal width, ventral tooth distinctly longer than dorsal tooth. Occipital carina meeting hy- postomal carina. Epomia short and weak. Sternaulus sharp and reaching middle coxa. Epicnemial carina curving posteri- orly and ending near subalarum. Hind margin of metanotum regular, but front margin of propodeum with strong tooth- like projections towards each side of postscutellum. Area between metanotum and propodeum moderately deep, wide in between tooth-like projections, narrow lat- erad of it, forming a polished smooth trough. Propodeum about 1.25 as long as wide. Propodeal spiracle large, elongate. Pleural carina strong, with cross ridges along its length. Propodeum entirely pol- ished smooth; both anterior and posterior transverse carinae conspicuous and com- plete; the anterior carina regular and somewhat acuminate, the posterior carina more projected centrally, laterally expand- ed into a somewhat translucent crest. Jux- tacoxal carina strong but small. Fore wing cell l + 2Rs about as long as width of pterostigma, about 1.15 wider Volume 14, Number 2, 2005 133 than higher, crossveins 2r-m and 3r-m ap- proximately parallel, about same length. Ramellus absent. Crossvein lcu-a distinct- ly but shortly basad of lM + Rs. Hind wing vein 1-Cu 1.3 length of crossvein cu-a; vein 2-1 A ending very near wing margin. Fore tibia in female distinctly swollen. Fourth segment of all tarsi moderately to deeply bilobed. First metasomal tergite at base with a lateral flange, without a tri- angular tooth; dorsolateral carina absent; ventro-lateral carina present, more distinct apically, beyond spiracle; spiracle at basal 0.47, strongly protuberant; sternite about 0.61 the length of tergite. Ovipositor 1.74 as long as hind tibia, projecting beyond metasoma for 5 times its own length; ovi- positor sheath about 1.44 as long as hind tibia. Lower valve of ovipositor with a dis- tinct subapical lobe that encloses most of upper valve; apex with 9 teeth. Etymology. — From the Greek dynos, meaning large, in reference to the body size, and informal reference to the Neo- tropical region, where the species was col- lected. Comments. — The genus runs to Ccstnis Townes in the key provided by Townes (1970) for the world genera of Gabuniina, but can be isolated from it by having a very short epomia (vs. long), propodeum fully smooth in front and behind both transverse carinae (vs. rugulose in be- tween carinae), petiole elongate, almost straight in lateral view (vs. short and dis- tinctly bent centrally), petiolar spiracle in lateral view fully dorsal (vs. centered), fore wing vein 2-Cu nearly of the same length of crossvein 2cu-a (vs. distinctly longer), areolet subquadrate, crossveins 2r-m and 3r-m nearly parallel (vs. distinct- ly pentagonal, 2r-m and 3r-m strongly convergent anteriorly), vein 2-M much longer than 3-M (vs. nearly of same length), and clypeus margin without a tooth (vs. usually with a tooth). Dincotropica lissa Aguiar, n. sp. Figs. 13-18 Description. — Female (Holotype). Fore wing 16.1 mm long. Clypeus and Mandi- ble as for the genus description. Occipital carina low and sharp throughout, joining the weakly raised hypostomal carina. Pronotum entirely polished smooth; epomia strong, sharp, short, transverse. Mesonotum: notauli anteriorly distinct, on posterior half weakly convergent and fused to dense longitudinal rugulosities, which end at scuto-scutellar groove; me- sonotum otherwise weakly alutaceous. Scutellum with micropunctures derived from weak pilosity. Mesepisternum pol- ished smooth; epicnemial carina ending near, but not at, subalarum; sternaulus strongly sinuous, sharp and narrow from epicnemial carina to base of hind coxa; without any indication of a depression be- tween sternaulus and speculum. Meta- pleuron mostly polished smooth, with weak transverse rugulosities on its poste- rior 0.25. Propodeum mostly polished smooth, scarcely pilose; spiracle large and elongate; anterior and posterior transverse carinae strong and complete; anterior ca- rina crossing shaft between propodeum and metanotum, ending on carinal trian- gle; posterior carina laterally somewhat raised, forming a crest, ending on pleural carina, which is strong, complete. Legs: all preapical tarsomeres deeply bilobed. Me- tasoma: first metasomal tergite, spiracle and sternite as in the genus description; Tl with a few punctures centrally, other- wise polished smooth; T2 with weak oblique creases; T3-8 polished smooth, densely covered by short golden pilosity. Ovipositor valves and sheath as in the ge- nus description. Wing venation as de- scribed for the genus. Color. — Head: scape yellow; pedicel, fla- gellomeres 1-2, and basal 0.9 of 3rd, black; apex of 3rd, 4-10 entirely, and most of 11"', yellow; 12"1 to apical entirely dark brown; 22 flagellomeres total, the apical one mod- erately compressed and protruded at apex. Orbital band yellow, complete ex- cept very briefly interrupted at bottom of eye; taking entire width of gena; narrow- ing dorsally to as narrow as 0.4 the dis- 134 Journal of Hymenoptera Research tance between eye margin and occipital carina at temple; face, clypeus, and la- brum entirely yellow; labial and maxillary palpi brownish; mandible basally orange, its teeth black. Mesosoma, including pro- podeum, reddish brown, darker dorsally. Legs reddish brown except yellowish to yellow fore to hind tibiae; tarsi darker from base to apex. Metasoma black; Tl dark red on basal 0.7, black apically, with yellow spot on central 0.5 of apical mar- gin; T2-6 black, with yellow stripe on api- cal and lateral margins, apical stripe inter- rupted laterally, not fused to lateral stripe; 17 black, with yellow stripe on apical and lateral margins, fused and continuous; T8 black with yellow stripe on ventral margin only. Ovipositor dark reddish brown; sheaths dark brown. Wings with brownish tint. Male. — Unknown. Material. — Holotype 9 DZUP "Ouro Preto, d'Oeste, RO, (12-I?}-1987, C. Elias, leg"; "Projeto Po- lonoroestc." Etymology. — From the Latin lissos, meaning polished, smooth; in reference to the body sculpture. Distribution record. — Northwestern Bra- zil (RO). KEY TO GENERA OF NEOTROPICAL GABUNIINA 1(0). Pleural carina present, even if incomplete 2 Pleural carina absent 5 2(1). Hind wing vein 1-Cu distinctly longer than crossvein cu-a; clypeal margin centrally normally with one or two small teeth 3 Hind wing vein 1-Cu nearly the same length or distinctly shorter than crossvein cu-a; clypeal margin centrally without a small tooth 4 3(2). Ovipositor just basad of apical teeth smooth; fore wing crossveins 2r-m and 3r-m more or less parallel; fore wing vein 2-Cu nearly as long as, or shorter than crossvein 2cu-a; hind wing vein M+Cu strongly convex Digonocryptus Viereck Ovipositor just basad of apical teeth with a distinctly microsculptured area; fore wing crossveins 2r-m and 3r-m distinctly convergent toward anterior margin of wing; fore wing vein 2-Cu distinctly longer than crossvein 2cu-a; hind wing vein M+Cu weakly convex Cestnts Townes 4(2). Posterior transverse carina of propodeum complete, strong; ovipositor just basad of apical teeth with distinctly microsculptured area; fore wing crossvein leu-a very nar- rowly basad of 1M+Rs; fore wing vein 2-M much longer than vein 3-M; fore wing vein 2Cu slightly longer than crossvein 2cu-a Dineotropica n. gen. Posterior transverse carina of propodeum absent; ovipositor just basad of apical teeth smooth; fore wing crossvein leu-a basad of lM + Rs by 0.3 its own length; fore wing vein 2-M distinctly shorter than vein 3-M; fore wing vein 2Cu much shorter than crossvein 2cu-a Prosthaporus Porter 5(1). Clypeal margin centrally without a small tooth; first metasomal tergite somewhat elon- gate, maximum length/ (maximum width— minimum width) about 4.2 6 Clypeal margin centrally with one or two small teeth; first metasomal tergite relatively short and triangular, maximum length/ (maximum width— minimum width) about 2.5-3.0 7 Vopodeum polished, unsculptured, entirely devoided of carinae; hind wing vein 1-Cu somewhat shorter than crossvein cu-a; fore wing crossvein leu-a basad of lM + Rs 36 its length; fore wing vein 2-Cu with about same length as crossvein 2cu-a; wing vein 2-M approximately as long as vein 3-M TrypHa Townes m granulose to granulose-striate, the anterior transverse carina complete, al- Volume 14, Number 2, 2005 135 most straight; hind wing vein 1-Cu distincly longer than crossvein cu-a; fore wing crossvein lcu-a narrowly basad or opposite vein lM + Rs; fore wing vein 2-Cu greatly shortened, so that 3-Cu almost meets 1-Cu; fore wing vein 2-M distinctly longer than vein 3-M Lagarosoma Gupta et Gupta 7(5). Fore wing cell l + 2Rs small, 0.2-0.3 as high as length of vein 2m-cu; fore wing vein 1-Rs+M with bulla placed centrally; fore wing hyaline or infuscate but never with dark bands 8 Fore wing cell l + 2Rs very large, 0.50-0.95 as high as length of vein 2m-cu; fore wing vein 1-Rs+M with bulla apical, reaching cell l+2Rs; fore wing hyaline, with three dark bands Nesolinoceras Ashmead 8(7). Fore wing crossveins 2r-m and 3r-m parallel or nearly so; hind wing vein 1-Cu with nearly the same length or slightly shorter than crossvein cu-a; fore wing vein 2-M substantially longer than vein 3-M Agonocryptus Cushman Fore wing crossveins 2r-m and 3r-m distinctly convergent towards anterior margin of wing; hind wing vein 1-Cu about twice as long as crossvein cu-a; fore wing vein 2-M approximately as long as vein 3-M Fenixia n. gen. ACKNOWLEDGEMENTS David B. Wahl contributed with valuable sugges- tions, guidance, and ideas to this work. Ian D. Gauld and Andrew Bennett also contributed on an earlier stage of the manuscript. Gavin R. Broad, Ilari E. Saaksjarvi, Donald L. J. Quicke and an anonymous reviewer read the submitted manuscript and re- turned very useful suggestions, corrections and in- formation. Vinalto Graf and Gabriel A. R. Melo loaned the specimens, after allowing the author to search the entire DZUP collection for unsorted Ga- buniina. Elisa Queiroz Garcia provided shelter and good company during the author's stay at DZUP. Lu- ciana Musetti, David Wahl, Santosh Gupta and An- drew Bennett helped with the literature. Rogerio Rosa da Silva kindly calculated the graph for Fig. 1. Fund- ing for this project was provided by FAPESP (Brazil), paid directly to the author (Processes 00/05704-6 and 03/08585-6), or acquired from specific funds allocated to this work in a connected project of C. Roberto F. Brandao (FAPESP process 98/05083-0). LITERATURE CITED Arnett, R. H., Jr., G. A. Samuelson, and G. M. Nish- ida. 1993. The Insect and Spider Collections of the World. Sandhill Crane Press, Gainesville, Florida. Broad, G. R. and D. L. J. Quicke. 2000. The adaptive significance of host location by vibrational soud- ing in parasitoid wasps. Proceedings of the Royal Society of London B 267: 2403-2409. Gauld, 1. D., D. B. Wahl, K. Bradshaw, P. Hanson, and S. Ward. 1997. The Ichneumonidae of Costa Rica, 2. Introduction and keys to species of the smaller subfamilies, Anomaloninae, Ctenopel- matinae, Diplazontinae, Lycorininae, Phrudinae, Tryphoninae (excluding Netelia) and Xoridinae, with an appendix on the Rhyssinae. Memoirs of the American Entomological Institute 57: 1-485. Goloboff, P. A. 1993a. Estimating character weights during tree search. Cladistics 9: 83-91. Goloboff, P. A. 1993b. NONA version 1.0. Published by the author, San Miguel de Tucuman, Argen- tina. Goloboff, P. A. 1997. PIWE version 2.6. Published by the author, San Miguel de Tucuman, Argentina. Gupta, S. 1983. A revision of the genus Prosthoporus (Hymenoptera: Ichneumonidae). Contributions of the American Entomological Institute 20: 245-253. Gupta, S. and V. K. Gupta. 1983. Ichneumonologia Orientalis, Part IX. The tribe Gabuniini (Hyme- noptera: Ichneumonidae). Oriental Insects Mono- graphs 10: 1-313. Gupta, V. K. and J. K. Jonathan. 1970. The genera of the Dagathia-complex (Hymenoptera: Ichneu- monidae). Oriental Insects 3: 389-393. Gupta, V. K. and J. K. Jonathan. 1971. A new Mesos- tenine genus Tanepomidos from India (Hymenop- tera: Ichneumonidae). Oriental Insects 5: 145-148. Henaut, A. 1990. Study of the sound produced by Pimpla instigator (Hymenoptera: Ichneumonidae) during host selection. Entomophaga 35: 127-139. Horstmann, K. 2001. Revision der von Johann Chris- tian Fabricius beschriebenen Ichneumonidae (Hymenoptera)- Beitriige zur Entoiuologie 51: 7-50. Laurenne, N. M., G. R. Broad, and D. L. Quicke. 2003 (2002). Preliminary molecular phvlogenetic analysis of Crytpinae and related taxa based on 26S D2 + D3 rDNA analysed using POY. Pp. 229-233 in: Melika, G. and C. Thuroczy, eds. Parasitic wasps: Evolution, Systematics, Biodiver- sity and Biological Control. Agroinform, Buda- pest, Hungary. Nixon, K. C. 1999a. The Parsimony Ratchet, a new 134 Journal of Hymenoptera Research tance between eye margin and occipital carina at temple; face, clypeus, and la- brum entirely yellow; labial and maxillary palpi brownish; mandible basally orange, its teeth black. Mesosoma, including pro- podeum, reddish brown, darker dorsally. Legs reddish brown except yellowish to yellow fore to hind tibiae; tarsi darker from base to apex. Metasoma black; Tl dark red on basal 0.7, black apically, with yellow spot on central 0.5 of apical mar- gin; T2-6 black, with yellow stripe on api- cal and lateral margins, apical stripe inter- rupted laterally, not fused to lateral stripe; T7 black, with yellow stripe on apical and lateral margins, fused and continuous; T8 black with yellow stripe on ventral margin only. Ovipositor dark reddish brown; sheaths dark brown. Wings with brownish tint. Male. — Unknown. Material— Holotype 9 DZUP "Ouro Preto, d'Oeste, RO, ( 12-171-1987, C. Elias, leg"; "Projeto Po- lonoroeste." Etymology. — From the Latin lissos, meaning polished, smooth; in reference to the body sculpture. Distribution record. — Northwestern Bra- zil (RO). KEY TO GENERA OF NEOTROPICAL GABUNIINA 1(0). Pleural carina present, even if incomplete 2 Pleural carina absent 5 2(1). Hind wing vein 1-Cu distinctly longer than crossvein cu-a; clypeal margin centrally normally with one or two small teeth 3 Hind wing vein 1-Cu nearly the same length or distinctly shorter than crossvein cu-a; clypeal margin centrally without a small tooth 4 3(2). Ovipositor just basad of apical teeth smooth; fore wing crossveins 2r-m and 3r-m more or less parallel; fore wing vein 2-Cu nearly as long as, or shorter than crossvein 2cu-a; hind wing vein M+Cu strongly convex Digonocryptus Viereck Ovipositor just basad of apical teeth with a distinctly microsculptured area; fore wing crossveins 2r-m and 3r-m distinctly convergent toward anterior margin of wing; fore wing vein 2-Cu distinctly longer than crossvein 2cu-a; hind wing vein M + Cu weakly convex Oestrus Townes 4(2). Posterior transverse carina of propodeum complete, strong; ovipositor just basad of apical teeth with distinctly microsculptured area; fore wing crossvein leu-a very nar- rowly basad of lM + Rs; fore wing vein 2-M much longer than vein 3-M; fore wing vein 2Cu slightly longer than crossvein 2cu-a Dineotropica n. gen. Posterior transverse carina of propodeum absent; ovipositor just basad of apical teeth smooth; fore wing crossvein leu-a basad of lM + Rs by 0.3 its own length; fore wing vein 2-M distinctly shorter than vein 3-M; fore wing vein 2Cu much shorter than crossvein 2cu-a Prosthoporus Porter 5(1). Clypeal margin centrally without a small tooth; first metasomal tergite somewhat elon- gate, maximum length/(maximum width— minimum width) about 4.2 6 Clypeal margin centrally with one or two small teeth; first metasomal tergite relatively short and triangular, maximum length/ (maximum width— minimum width) about 2.5-3.0 7 Propodeum polished, unsculptured, entirely devoided of carinae; hind wing vein 1-Cu somewhat shorter than crossvein cu-a; fore wing crossvein leu-a basad of lM + Rs 0.36 its length; fore wing vein 2-Cu with about same length as crossvein 2cu-a; tore wing vein 2-M approximately as long as vein 3-M Tn/pha Townes ■opodeum granulose to granulose-striate, the anterior transverse carina complete, al- Volume 14, Number 2, 2005 135 most straight; hind wing vein 1-Cu distincly longer than crossvein cu-a; fore wing crossvein lcu-a narrowly basad or opposite vein lM + Rs; fore wing vein 2-Cu greatly shortened, so that 3-Cu almost meets 1-Cu; fore wing vein 2-M distinctly longer than vein 3-M Lagarosoma Gupta et Gupta 7(5). Fore wing cell l + 2Rs small, 0.2-0.3 as high as length of vein 2m-cu; fore wing vein 1-Rs+M with bulla placed centrally; fore wing hyaline or infuscate but never with dark bands 8 Fore wing cell l+2Rs very large, 0.50-0.95 as high as length of vein 2m-cu; fore wing vein 1-Rs+M with bulla apical, reaching cell l+2Rs; fore wing hyaline, with three dark bands Nesolinoceras Ashmead 8(7). Fore wing crossveins 2r-m and 3r-m parallel or nearly so; hind wing vein 1-Cu with nearly the same length or slightly shorter than crossvein cu-a; fore wing vein 2-M substantially longer than vein 3-M Agonocryptus Cushman Fore wing crossveins 2r-m and 3r-m distinctly convergent towards anterior margin of wing; hind wing vein 1-Cu about twice as long as crossvein cu-a; fore wing vein 2-M approximately as long as vein 3-M Fenixia n. gen. ACKNOWLEDGEMENTS David B. Wahl contributed with valuable sugges- tions, guidance, and ideas to this work. Ian D. Gauld and Andrew Bennett also contributed on an earlier stage of the manuscript. Gavin R. Broad, Ilari E. Saaksjarvi, Donald L. J. Quicke and an anonymous reviewer read the submitted manuscript and re- turned very useful suggestions, corrections and in- formation. Vinalto Graf and Gabriel A. R. Melo loaned the specimens, after allowing the author to search the entire DZUP collection for unsorted Ga- buniina. Elisa Queiroz Garcia provided shelter and good company during the author's stay at DZUP. Lu- ciana Musetti, David Wahl, Santosh Gupta and An- drew Bennett helped with the literature. Rogerio Rosa da Silva kindly calculated the graph for Fig. 1. Fund- ing for this project was provided by FAPESP (Brazil), paid directly to the author (Processes 00/05704-6 and 03/08585-6), or acquired from specific funds allocated to this work in a connected project of C. Roberto F. Brandao (FAPESP process 98/05083-0). LITERATURE CITED Arnett, R. H., Jr., G. A. Samuelson, and G. M. Nish- ida. 1993. The bisect and Spider Collections of the World. Sandhill Crane Press, Gainesville, Florida. Broad, G. R. and D. L. J. Quicke. 2000. The adaptive significance of host location by vibrational soud- ing in parasitoid wasps. Proceedings of the Royal Society of London B 267: 2403-2409. Gauld, I. D., D. B. Wahl, K. Bradshaw, P. Hanson, and S. Ward. 1997. The Ichneumonidae of Costa Rica, 2. Introduction and keys to species of the smaller subfamilies, Anomaloninae, Ctenopel- matinae, Diplazontinae, Lvcorininae, Phrudinae, Tryphoninae (excluding Netelia) and Xoridinae, with an appendix on the Rhyssinae. Memoirs of the American Entomological Institute 57: 1^485. Goloboff, P. A. 1993a. Estimating character weights during tree search. Cladistics 9: 83-91. Goloboff, P. A. 1993b. NONA version 1.0. Published by the author, San Miguel de Tucuman, Argen- tina. Goloboff, P. A. 1997. PIWE version 2.6. Published by the author, San Miguel de Tucuman, Argentina. Gupta, S. 1983. A revision of the genus Prosthoporus (Hymenoptera: Ichneumonidae). Contributions of the American Entomological Institute 20: 245-253. Gupta, S. and V. K. Gupta. 1983. Ichneumonologia Orientalis, Part IX. The tribe Gabuniini (Hyme- noptera: Ichneumonidae). Oriental Insects Mono- graphs 10: 1-313. Gupta, V. K. and J. K. Jonathan. 1970. The genera of the Dagathia-comp\ex (Hymenoptera: Ichneu- monidae). Oriental Insects 3: 389-393. Gupta, V. K. and J. K. Jonathan. 1971. A new Mesos- tenine genus Tanepomidos from India (Hymenop- tera: Ichneumonidae). Oriental Insects 5: 145-148. Henaut, A. 1990. Study of the sound produced by Pimpla instigator (Hymenoptera: Ichneumonidae) during host selection. Etitomophaga 35: 127-139. Horstmann, K. 2001. Revision der von Johann Chris- tian Fabricius beschriebenen Ichneumonidae (Hymenoptera). Bcitragc zur Entomologie 51: 7-50. Laurenne, N. M., G. R. Broad, and D. L. Quicke. 2003 (2002). Preliminary molecular phylogenetic analysis of Crytpinae and related taxa based on 26S D2 + D3 rDNA analysed using POY. Pp. 229-233 in: Melika, G. and C. Thuroczy, eds. Parasitic wasps: Evolution, Systematics. Biodiver- sity and Biological Control. Agroinform, Buda- pest, Hungary. Nixon, K. C. 1999a. The Parsimony Ratchet, a new 136 Journal of Hymenoptera Research method for rapid parsimony analysis. Cladistics 15: 407-414. Nixon, K. C. 1999b. Winclada (BETA) ver. 0.9.9. Pub- lished by the author, ITHACA, New York. Otten, H., F. L. Wackers, M. Battini, and S. Dorn. 2000. Efficiency of vibrational sounding in the parasitoid Pimpla turionellae is affected by female size. Animal Behaviour 61: 671-677. Otten, H., F. L. Wackers, N. Isidoro, R. Romani and S. Dorn. 2003 (2002). The subgenual organ in Pimpla turionellae L. (Hymenoptera Ichneumoni- dae): ultrastructure and behavioral evidence for its involvement in vibrational sounding. Redia 85: 61-76. Townes, H. K. 1970. The genera of Ichneumonidae. Part 2. Memoirs of the American Entomological In- stitute 12: 1-537. Townes, H. and M. Townes. 1962. Ichneumon-flies of America north of Mexico: 3. Subfamily Gelinae, tribe Mesostenini. United States National Museum Bulletin 216: 1-602 + i-viii. Venables, W. N. and B. D. Ripley. 1997. Modern ap- plied statisties with S-PLUS. Second edition. Springer- Verlag, New York. 548 pp. Vilhelmsen, L., N. Isidoro, F. Bin, H. H. Basibuyuk, and D. L. J. Quicke. 2001. Host location and ovi- position in a basal group of parasitic wasps: the subgenual organ, ovipositor apparatus and as- sociated structures in the Orussidae (Hymenop- tera, Insecta). Zoomorphology 121: 63-84. Yu, D. S. and K. Horstmann. 1997. A catalogue of world Ichneumonidae (Hymenoptera). Memoirs of the American Entomological Institute 58: 1-1558 + i-vi., 2 vols. J. HYM. RES. Vol. 14(2), 2005, pp. 137-150 Taxonomy, Biology, and Distribution of Seed Harvesting Ants in the Pheidole californica Complex (Hymenoptera: Formicidae) Dylan O. Burge Department of Biology, Duke University, Box 90338, Durham, NC 27708 email: dob7@duke.edu Abstract. — The Pheidole californica complex comprises three closely related and partly sympatric species of seed harvesting ants restricted to the western Nearctic region. An analysis of morpho- logical variation in the minor workers and major workers (soldiers) reveals that these species are more variable than previously thought. Nevertheless, morphological characteristics were uncov- ered that are consistently diagnostic for each species. Pheidole clementensis Gregg may be distin- guished from both Pheidole californica Mayr and Pheidole creightoni Gregg by the diagonal rather than longitudinal rugulae between the clypeus and the eye in lateral view. The closely related P. californica and P. creightoni are most readily distinguished from one another by the angle of their lateral cephalic setae. Pheidole californica has decumbent setae forming an angle of forty five de- grees or less with the lateral margins of the head in full face view, while the cephalic setae of P. creightoni emerge at an angle of approximately ninety degrees. Pheidole californica is broadly dis- tributed in the western Nearctic region. Pheidole creightoni and P. clementensis are more limited in their range, the former being found in foothill or plateau habitats in northern California, Oregon, Washington, and Nevada, while the latter is limited to foothill and coastal habitats of southern California and northern Baja California. The cosmopolitan genus Pheidole is rep- resented in the New World by more than 600 described species, which form a di- verse and ecologically important compo- nent of many ant communities. Perhaps as a result of their dominance and diversity, the New World Pheidole have received considerable taxonomic attention (e.g., Mayr 1870, Emery 1895, Creighton 1950, Gregg 1959), culminating in a recent monograph (Wilson 2003) that more than doubled the number of described species. Despite this attention the taxonomy of the New World Pheidole is not fully re- solved. Many taxa are refractory to taxo- nomic characterization because of similar- ity between the species, as well as exten- sive intraspecific variation both within and among populations. Pheidole californi- ca, P. creightoni, and P. clementensis com- prise one such assemblage, here termed the californica complex, which is part of the larger "pilifera group" of Wilson (2003). Among the three species included in this study, P. californica has both the larg- est geographic range and highest degree of intraspecific variation. In an attempt to encompass this variation, taxonomists of the late nineteenth and early twentieth century described additional species, and a number of subspecies and varieties (Em- ery 1895, Wheeler 1915, Cole 1933, 1936), although subsequent taxonomic work ac- knowledged the synonymy of most of these with P. californica (Creighton 1950, Gregg 1959, Wheeler and Wheeler 1986, Wilson 2003). In spite of all this taxonomic attention P. californica remains a difficult ant to char- acterize, and the precise boundary be- tween it and the other two species includ- ed in the study has never been rigorously examined. The purpose of the present con- tribution is to revise the taxonomy of the californica complex and to characterize 138 Journal of Hymenoptera Research each species in a way that takes into ac- count both intra- and interspecific varia- tion. MATERIALS AND METHODS At the beginning of this study I traveled to sixteen localities in northern California, Washington, and western Nevada to in- tensively sample the californica complex. Between three and twelve nest series were obtained from each species at a given lo- cality, along a transect that varied in length from one to twelve kilometers. The purpose of this sampling was to obtain baseline data on variation at the level of individual ants, colonies, populations, and species. In addition to the material obtained spe- cifically for this study, specimens were ex- amined from the collections of Philip S. Ward (PSWC), the Bohart Museum of En- tomology, University of California at Da- vis (UCDC), and the Los Angeles County Museum (LACM). Measurements of specimens for use in the diagnoses and bivariate plots were taken at 50 X using a Wild M5A micro- scope and a Nikon stage micrometer, and are presented to two decimal places. The following measurements and indices were utilized in this study: HL Head length: length of the head in full face view, measured from the anterior extremity of the clypeal margin to the midpoint of an imag- inary line drawn across the poste- rior margin of the head (after Ward 2000). HW Head width: maximum width of head in full face (frontal) view, not including the eyes. CI Cephalic Index: HW/HL. EL Eye length: maximum diameter of the eye, measured with the head in lateral view. REL Relative eye length: EL/HL. lotum width: maximum width pronotum, in dorsal view. DML Dorsal mesosomal length: length of the mesosoma measured in dorsal view, from the anterior extremity of the pronotum near the articulation with the head to the posterior ex- tremity of the propodeum (usually a cuticular flange near the articula- tion of the petiole). For this mea- surement, the specimen is adjusted in dorsal view so that the extremi- ties of the pronotum and propo- deum are simultaneously in focus. PPW Postpetiolar width: maximum width of postpetiole in dorsal view. SPECIES ACCOUNTS The three species included in this study are members of the "pilifera group", a pre- sumably monophyletic set of forty eight species of Pheidole, found almost exclu- sively in the Nearctic region, including Mexico (Wilson 2003). The group is char- acterized by reduced hypostomal teeth in the major, large eyes in the minor, and by the quadrate shape of the head in both major and minor (Wilson 2003). Within the pilifera group Wilson (2003) recognizes a cluster of species that he calls the "pili- fera complex," which includes the species P. calens, californica, carrolii, cavigenis, cle- mentensis, creightoni, hoplitica, littoralis, mi- cula, polymorpha, rugulosa, soritis, tepicana, and torosa. Although it is unclear whether these species represent a monophyletic group, they are united by the traits (in the major) of extensive cephalic sculpture and, except in P. carolli and P. littoralis, trans- verse rugulae or carinulae on the posterior cephalic vertex ("occipital lobes" in Wil- son 2003). Within this cluster of species I recognize a group of three species, here termed the californica complex, united by the trait of highly developed transverse ru- gulae (see Wilson 2003: 23) on the cephalic dorsum of the major. In 1915, W.M. Wheeler described P. cal- ifornica subsp. micula. Gregg (1959) later recognized this as a distinct species, P. mi- cula. Although I have not examined type Volume 14, Number 2, 2005 139 material for this species, I have concluded that P. micula is not a member of the cali- fornica complex, because it lacks highly de- veloped transverse rugulae on the cephal- ic dorsum of the majors. In his recent monograph, E. O. Wilson (2003) describes the dorsal cephalic sculpture of P. micula majors as consisting of transverse carinulae that sometimes wrap downward at the sides of the head. Wilson also presents the caveat, in the form of a personal commu- nication from Stefan Cover, that not all specimens of P. micula possess such exten- sive cephalic sculpture. Pheidole californica, P. clementensis, and P. creightoni are unique among the members of the pilifera-group in having majors with highly developed ce- phalic sculpture consisting of transverse rugulae, as opposed to carinulae (see Wil- son 2003: 23), which cover the posterior cephalic vertex, extend at least one fourth of the way to the clypeus in full face view, and are at least partly visible in lateral view. Thus, I am leaving P. micula out of the californica complex. Nevertheless, the question of a possible affinity of P. micula with members of the californica complex warrants further examination. The diagnoses presented in the follow- ing species accounts are a summary of morphological features that are usually characteristic of the species and therefore useful in identification. The features that are most consistently diagnostic of species are presented in bold face. Intraspecific variation in the remaining traits makes them less useful for identification. In ad- dition, the illustrations (Figs. 5-21) should not be interpreted as comprehensive rep- resentations of the respective species. Ce- phalic sculpture and setae vary dramati- cally within these species, and most of the differences shown in these figures are not diagnostic for the species. Reliably diag- nostic features are indicated in the figures with arrows. Pheidole californica Mayr 1870 (Figs. 5-8, 21) Pheidole californica Mayr 1870:987. Description of worker. Pheidole oregonica Emery 1895:291. Synonymy by Wilson (2003:564). Pheidole californica var. incenata Wheeler 1915: 407. Synonymy by Creighton (1950:173). Pheidole californica var. satura Wheeler 1915:407. Synonymy by Creighton (1950:173). Pheidole californica subsp. nevadensis Wheeler 1915:408. Synonymy by Wheeler and Wheel- er (1986:13). Pheidole californica subsp. pyramidensis Emery 1922:105. Replacement name for subsp. nev- adensis (junior primary homonym of P. pubi- ventris var. nevadensis Forel 1901:353). Syn- onymy by Wheeler and Wheeler (1986:13). Pheidole californica var. shoshoni Cole 1933: 618. Synonymy by Gregg (1959:19). Pheidole californica var. hagermani Cole 1936: 35. Synonymy by Creighton (1950:173). Pheidole californica Mayr; Wheeler and Wheeler 1972:243 (description of larva). Major worker measurements (n = 65): HL 0.98-1.28, HW 0.86-1.21, CI 0.87-0.97, EL 0.14-0.22, REL 0.12-0.19, PrW 0.40- 0.56, DML 0.83-1.07, PPW 0.17-0.31. Diagnosis of major worker. — Head in full face view subquadrate. Lateral margins of head weakly to strongly convex. Lobes of posterior cephalic vertex weakly devel- oped, and the notch of the vertex therefore shallow (Fig. 6); rugulae originating on the lateral clypeal margin straight and longi- tudinal; those rugulae originating on the part of the clypeus directly in front of the eye terminating abruptly at the eye (Fig. 5); eyes usually large in relation to the length of the head (Fig. 5); in full face view, setae emerging laterally from the head decumbent, forming an angle of for- ty five degrees or less with the integument (Fig. 6); propodeal spines in side view typ- ically in the form of equilateral triangles produced from the lateral margination of the propodeum (Fig. 7). Minor worker measurements (n = 20): HL 0.53-0.72, HW 0.47-0.66, CI 0.86-0.94, EL 0.12-0.16, REL 0.21-0.26, PrW 0.30-0.39, DML 0.67-0.79, PPW 0.12-0.18. Diagnosis of minor worker. — Mesosoma elongate (DML 0.67-0.79); foveolate (see Wilson 2003: 22) sculpturing dense and 142 Journal of Hymenoptera Research V. Chamberton); UC Santa Cruz campus, 230m (P. S. Ward); Shasta Co.: 1km WNW Lamoine, 630m (P. S. Ward); Enterprise, 175m (P. S. Ward); Siskiyou Co.: 17km SW Scott Bar, 640m (P. S. Ward); 26.8km S Tule Lake, 1420m (D. O. Burge); Weed (A. C. Cole); Solano Co.: 10km NE Dixon, 15m (P. S. Ward); 13km NW Dixon, 35m (P. S. Ward); 13km NW Rockville, 85m (P. S. Ward); Allendale (R. Waegell); Cold Canyon, 19km NNW Vacaville, 107m (D. O. Burge); Cold Can- yon, 19km NNW Vacaville, 120m (P. S. Ward); Cold Canyon, 19km NNW Vacaville, 120m (D. M. Olson) Cold Canyon, 19km NNW Vacaville, 600m (P. S Ward); Jepson Prairie, 18km S Dixon, 10m (P. S Ward); South end of Lake Solano, 40m (R. Waegell) Sonoma Co.: 1km NNE Sonoma, 170m (P. S. Ward); 2km ENE Glen Ellen, 150m (P. S. Ward); 6km N Son- oma, 300m (P. S. Ward); Pepperwood Ranch, 15km N Santa Rosa, 360m (P. S. Ward); Sonoma, 25m (P. S. Ward); Stanislaus Co.: 22km WSW Patterson, 350m (D. O. Burge); Del Puerto Canyon, 22km WSW Patterson, 350m (P. S. Ward); Del Puerto Canyon, Frank Raines Regional Park, 340m (A. L. Wild); Sutter Co.: 7km NNW Sutter, 210m (P. S. Ward); Tehama Co.: 6km E Payne's Creek, 720m (P. S. Ward); 8.02km SSE Paynes Creek, 308m (D. O. Burge); 17.3km ESE Redbluff, 352m (D. O. Burge); hwy. 36, 5.8mi NE jet. w/hwy 99 (R. R. Snelling and P. Mehlhop); Redding (A. C Cole); Tuolumne Co.: 3km SW Cold Springs, 1700m (P. S. Ward); 2mi W Chinese Camp (J. I. Stage) Sweetwater Campground, Stanislaus N.F. (S. Bloem) Ventura Co.: 29km WNW Stauffer, 1460m (P. S Ward); Yolo Co.: 2.2km WSW Davis, 18m (D. O Burge); 2km SSE Dobbins, 560m (P. S. Ward); 10 km N Davis, 10m (P. S. Ward); 10km W Winters, 60m (P. S. Ward); 13km W Rumsey, 710m (P. S. Ward); Ber- ryessa Peak, 930m (P. S. Ward); Davis (P. S. Ward); Grasslands Regional Park, 8km SE Davis, 10m (P. S. Ward); Yuba Co.: Sierra Foothill Range, 18mi NE Mar- ysville (F. A. Ludtke). Idaho: Elmore Co.: 5.5mi E Hammett (R. R. Snelling); Gooding Co.: Hagerman (A. C. Cole); Oivyhee Co.: 3.3mi S Given's Hot Springs (R. R. Snelling); Twin Falls Co.: Twin Falls (A. C. Cole). Nevada: Humboldt Co.: Calico Mtns., 35mi N Gerlach (W. S. Creighton); Washoe Co.: 5km S Nixon, 1185m (P. S. Ward); 8.67km W Nixon, 1140m (D. O. Burge); Hanging Rock Canyon, 1750m (P. S. Ward); S end Pyramid Lake, 1140m (P. S. Ward). Oregon: Deschutes Co.: Smith Rock State Park, 880m (P. S. Ward); Jose- phine Co.: 8km SSW Cave Junction, 430m (P. S. Ward); lake Co.: 5mi S Plush (R. R. Snelling); Wasco Co.: Mau- pirt (W. S. Creighton). Utah: Salt Lake Co.: Salt Lake City (Grundmann). Washington: Adams Co.: Mc- Mannanuin Lake. 260m (P. S. Ward); Grant Co.: rtctvman< 260m (P. S. Ward); King Co.: Seat- Kin, m Co.: Riverside, 290m (D. O. I' iwai (VV. M. Mann). Pheidole creightoni Gregg 1955 (Figs. 13-16) Pheidole creightoni Gregg 1955: 19 (w, q, m). Major worker measurements (n = 34): HL 1.15-1.37, HW 1.05-1.30, CI 0.88-0.96, EL 0.17-0.21, REL 0.13-0.17, PrW 0.44-0.56, DML 0.94-1.10, PPW 0.25-0.42. Diagnosis of major worker. — Head su- bquadrate in full face view (Fig. 14); lobes of the anterior cephalic vertex developed, so that in full face view the medial notch is pronounced (Fig. 14); rugulae that orig- inate on the lateral clypeal margin straight and longitudinal; those rugulae originat- ing on the part of the clypeus directly in front of the eye terminate abruptly at the eye (Fig. 13); eyes small in relation to the length of the head (REL 0.13-0.17; Fig. 13); due to the development of the lobes of the posterior cephalic vertex, eyes appear to be placed near the posterior clypeal mar- gin, often within the first third or fourth of the length of the head in lateral view (Fig. 13); pilosity short and erect; in full face view, setae emerging laterally from the head erect, forming an angle of ap- proximately ninety degrees with the integ- ument (Fig. 14); propodeal spines in side view generally in the form of long blunt pegs (Fig. 15); lateral margination of the propodeum not usually well developed. Minor worker measurements (n = 8): HL 0.53-0.62, HW 0.48-0.57, CI 0.89-0.93, EL 0.13-0.15, REL 0.22-0.26, PrW 0.31-0.35, DML 0.67-0.75, PPW 0.14-0.17. Diagnosis of the minor worker. — Mesosoma elongate (DML 0.67-0.75); foveolate sculp- turing patchy on the mesosoma in side view, interspersed on the metapleuron, mesopleuron, and propodeum with patch- es of smooth, shining integument; propo- deal spines normally sharp and elongate (Fig. 16); lateral margination of the propo- deum usually weakly developed. Comments. — At five localities P. creigh- toni and P. californica are known to co-oc- cur (Fig. 24). At four of these sites the spe- cies appear to be distinct, but at Dye Volume 14, Number 2, 2005 143 Creek, California, a fraction of the colonies (2 out of 9) sampled along a three kilo- meter transect yielded workers that were difficult to assign to either species. The morphology of these ants is curiously in- termediate between P. California! and P. creightoni, "bridging the gap" between the two species. In general, the unusual colo- nies yielded workers with the size and col- oration of P. creightoni, but with the de- cumbent lateral cephalic setae that are di- agnostic of P. californica. In general, ants from the two unusual colonies resemble collections of P. californica (three out of nine colonies along the transect) much more than P. creightoni (four out of nine colonies along the transect), and they were treated as P. californica for the analysis of morphometric data. The presence of typical P. californica and P. creightoni at Dye Creek points to hy- bridization as a possible explanation for the existence of morphologically interme- diate colonies of Pheidole at that site. With- in northern California, the range of P. cal- ifornica appears to extend from the north- ern Sacramento Valley into the surround- ing foothills and plateau areas, while the range of P. creightoni appears to extend from the foothill and plateau regions to the edge of the Sacramento Valley (see Fig. 24). It may be that that the Dye Creek region forms part of a hybrid zone be- tween the two species. Resolution of this problem will probably require both genet- ic analysis and more intensive sampling in the foothills area of northern California. Material examined (PSWC, UCDC, LACM). UNITED STATES California: Mendocino Co.: 4.59km NNE Hopland, 292m (D. O. Burge); 7.7km NNE Hopland, 825m (D. O. Burge); Hopland Field Station, 240m (P. S. Ward); Siskiyou Co.: 1.5mi NE Ga- zelle, 2600' (R. R. Snelling); Weed (A. C. Cole); Te- hama Co.: 17.3km ESE Redbluff, 352m (D. O. Burge); 34.4km N Forest Ranch, 756m (D. O. Burge); Trinity Co.: Weaverville, Democrat Gulch (D. M. Gordon). Nevada: Lander Co.: Battle Mm. (A. C. Cole); Hum- boldt Co.: Winemucca (A. C. Cole); Lyon Co.: Fort Churchill State Historic Park, 1280m (P. S. Ward); Washoe Co.: 8.67km W Nixon, 1140m (D. O. Burge); S end Pyramid Lake, 1140m (P. S. Ward). Oregon: Jack- son Co.: 3mi E Applegate (W. S. Creighton); Josephine Co.: 8km SSW Cave Junction, 430m (P. S. Ward). Washington: Grant Co.: Corfu (C. H. Lavers). Pheidole clementensis Gregg 1969 (Figs. 17-20) Pheidole clementensis Gregg 1969: 93 (w). Major worker measurements (n = 14): HL 1.09-1.37, HW 0.94-1.23, CI 0.85-0.90, EL 0.14-0.18, REL 0.11-0.14, PrW 0.47-0.59, DML 0.92-1.11, PPW 0.23-0.34. Diagnosis of major worker. — Head usually rectangular (subquadrate and elongate) in full face view, lateral margins parallel and only mildly convex (Fig. 18); transverse rugulae near the vertex of the head ex- tremely coarse, such that in side view the head usually appears to have "brow ridg- es" (Fig. 17); in lateral view, rugulae orig- inating on the part of the clypeus directly anterior to the eye arranged diagonally (rather than longitudinally), running to- wards the ventral surface of the head, be- low the eye (Fig. 17); eyes small relative to the length of the head (REL 0.11-0.14; Fig. 17); eyes located near the posterior clypeal margin, often within the first third or fourth of the length of the head in side view (Fig. 17); propodeal spines in side view generally in the form of long blunt pegs (Fig. 19); lateral margination of the propodeum usually weakly developed. Minor worker measurements (n = 6): HL 0.52-0.61, HW 0.47-0.55, CI 0.89-0.92, EL 0.11-0.14, REL 0.19-0.22, PrW 0.30-0.35, DML 0.62-0.73, PPW 0.11-0.14. Diagnosis of minor worker. — Mesosoma short (DML 0.62-0.73); foveolate sculptur- ing on the lateral mesosoma nearly non-ex- istent, replaced by small, indistinct patches of sculpture mixed with more extensive ar- eas of smooth and shining integument; pro- podeal spines short, thin, and sharp-tipped (Fig. 20); lateral margination of the propo- deum usually weakly developed. Comments. — Majors of P. clementensis may be distinguished from majors of P. 144 Journal of Hymenoptera Research j^\ 1.0 mm H I Figs. 5-16. Pheidole major workers, full face view of head (6, 10, 14), lateral view of head (5, 9, 13), and lateral view of mesosoma (7, 11, 15). Pheidole minor workers, lateral view of mesosoma (8, 12, 16). Note that sculp- turing and setae of the antennal scapes and mesosoma have been omitted for clarity. 5-8: P. californica, 16.4 E Chico, California; 9-12: P. creightoni-like variant of P. californica, 4.59 km NNE Hopland, California; (-16: /'. crcl;<, ^>4.4 km N Forest Ranch, California. Volume 14, Number 2, 2005 145 1.0 mm 21a Figs. 17-21. Pheidole clementensis major worker, full face view of head (18), lateral view of head (17), and lateral view of mesosoma (19). Minor worker, lateral view of mesosoma (20). Pheidole californica major workers, full face view of head with sculpture, pilosity, and antennae omitted (21a-f). 17-20: /'. clementensis, Meling Ranch, Baja California, Mexico. 21a-c, three specimens of the P. creightoni-]ike variant of /'. California!, from a single colony; 21d-f, three specimens of P. californica sensu stricto from a single colony. 21 a-f: 19km NNW Vacaville, California. 146 Journal of Hymenoptera Research californica and P. creightoni by the angle of the rugulae between the eye and the clyp- eus in side view (see diagnoses). Pheidole clementensis overlaps more broadly with P. californica than P. creightoni for the metric measurements that were taken (see mea- surements in diagnoses), and bivariate plots tend to separate P. creightoni from P. clementensis more readily than they sepa- rate P. californica from P. clementensis (compare Figs. 1 and 2). The range of P. clementensis overlaps broadly with the range of P. californica (Figs. 22-23), arid the two species are sympatric at several sites. Although collections of the species from these sympatric zones are few, no intermediate forms are known, and hy- bridization does not appear to occur. The known range of P. clementensis is far re- moved from that of P. creightoni (Fig. 23). Material examined (PSWC, UCDC, LACM). MEXICO: Baja California: 8mi E El Rosario bridge (R. A. Johnson); Meling Ranch, 2450' (R. A. Johnson); Sierra San Borja 9.9mi N San Borja, 1970' (R. A. John- son). UNITED STATES: California: Los Angeles Co.: San Clemente Is., Pyramid Head (R. R. Snelling); Orange Co.: lmi NW El Toro, 450' (R. J. Hamton); Riverside Co.: Lake Skinner, 462m (A. V. Suarez); Skinner Res- ervoir, 4470' (T. Prentice); San Diego Co.: Camp Pen- dleton (J. H. Hunt); Chula Vista (E end), 160m (P. S. Ward); Miramar Naval Air Station (T. Prentice). IDENTIFICATION The following short key may be used to differentiate major workers of the three species of the californica complex. All spe- cies have highly developed cephalic sculp- ture consisting of transverse rugulae that cover the posterior cephalic vertex, extend at least one fourth of the way to the clyp- eus in full face view, and are at least part- ly visible in lateral view. 1. Rugulae originating on lateral clypeal margin (directly anterior to eye) straight and lon- gitudinal, terminating abruptly at the eye (Figs. 5, 9, 13) 2 - Rugulae originating on lateral clypeal margin (directly anterior to eye) arranged diagonally rather than longitudinally, running beneath the eye (Fig. 17) P. clementensis Gregg 2. In full face view, setae emerging laterally from the head decumbent, forming an angle of forty five degrees or less with the integument (Fig. 6) P. californica Mayr - In full face view, setae emerging laterally from the head erect, forming an angle of ap- proximately ninety degrees with the integument (Fig. 14) P. creightoni Gregg Gregg's (1959) key to the Pheidole of North America is also useful for identify- ing members of the californica complex. In light of newly discovered variation in crit- ical traits, however, some couplets of the key must be modified. Couplets 23 and 27 refer to the development of the post peti- olar connules, a trait that varies consider- ably within P. californica and P. creightoni. In general, P. creightoni has more highly de- veloped postpetiolar connules than P. cali- fornica, but the difference is not diagnostic. Couplet 27 of Gregg's key should be sim- plified to the following in order to prevent misidentification of P. creightoni that do not have well-developed postpetiolar connules. !7. Rugae on cephalic vertices of the major straight or wavy, but not reticulate; lateral post- petiolar connules usually very prominent 28 igae on cephalic vertices of the major notably reticulate and often coarse, but not retic- ilate; lateral postpetiolar connules usually blunt 29 Volume 14, Number 2, 2005 147 Couplet 24, which is based on the head fied in order to encompass newly discov- length (HL) of the major, must be modi- ered variation in this trait. 24. Head of major 0.85mm in length, or less 25 Head of major 0.98mm in length, or more 26 In their original form, couplets 33 and 34 of Gregg's key differentiate four subspecies of P. California!. As these subspecies are in synonymy with P. California!, I have pro- vided a replacement for couplets 33 and 34 that now includes P. clementensis, which was described subsequent to the publica- tion of Gregg's (1959) key. Although Gregg's (1969) description of P. clementensis includes a modification of the 1959 key that is intended to separate the new species from P. califoniica, the "brow ridges" to which Gregg's couplet refers are not a con- sistent characteristic of the species. Some specimens of P. californica have similar sculpture, and specimens of P. clementensis without "brow ridges" are known from southern California. The following couplet (adapted from the short key above) should replace Gregg's couplets 33 and 34. 33. Rugulae originating on lateral clypeal margin (directly anterior to eye) arranged diago- nally rather than longitudinally, running beneath the eye (Fig. 17) P. clementensis Gregg Rugulae originating on lateral clypeal margin (directly anterior to eye) straight and lon- gitudinal, terminating abruptly at the eye (Fig. 5) P. californica Mayr BIOLOGY AND DISTRIBUTION and Baja California (Fig. 22). Pheidole cali- rn. i_- i • i i_ i j i • fornica readily colonizes disturbed habitats I he biological observations included in J / .1 c ii • ,u i.u l such as road verges, fallow fields, and va the following synopses are those that ap- & pear to be characteristic of the species. Nevertheless, these traits vary consider- ably within each species. Distribution data are based on my own collections and in- formation associated with specimens that I examined in existing collections, which means that they are biased towards acces- sible areas. Thus, gaps on each map may indicate a lack of field work within a par- ticular region rather than a true gap in the distribution of the species. Pheidole californica. — This species is found in the greatest diversity of habitats, cant lots. Nest entrances are usually small and inconspicuous, making them difficult to find except when they are located in patches of bare soil. Midden piles are ac- cumulated, but this species does not usu- ally collect chaff around the nest in the manner of P. creightoni. Foraging usually takes place in the evening and during the night. Foraging is conducted primarily by minor workers. Nuptial flights tend to oc- cur in the evening and early night at most localities. In Davis, California, nuptial flights at a specific group of colonies were from valley grasslands to higher elevation observed to begin in early May, and con- mountainous or plateau areas (5 to 1750m, tinue sporadically until late June, mean = 600m) in Arizona, California, Ida- Pheidole creightoni. — This species is ho, Nevada, Oregon, Utah, Washington, found in foothill, mountain, and plateau 148 Journal of Hymenoptera Research a P. clementensis ■ P. crcightoni Figs. 22-23. Known distributions in the western United States and northern Mexico of Pheidole californica (22), and P. creightoni and P. clementensis (23). areas of California, Oregon, Washington, and Nevada (Fig. 23) at a higher average elevation than P. californica (300 to 1300m, mean = 725m). Pheidole creightoni is often found in less disturbed habitats than P. californica. Nest entrances are conspicuous, and usually located within tufts of grass or at the edge of partially buried stones. Significant amounts of discarded seed chaff are usually associated with nests, forming a soft "crater" around the en- trance, or a mound adjacent to the nest. The foraging habits of P. creightoni are similar to those of P. californica, although foraging columns of the former tend to contain a higher proportion of major workers. At several study sites where P. creightoni co-occurs with the seed harvesting ant Messor andrei (Mayr) these species tend to nest in close proximity. This phenomenon was first noted by W. S. Creighton, who originally collected P. creightoni in south- ern Oregon (Gregg 1955). At both Dye Creek and Hopland Research and Exten- sion Center, California, P. creightoni colo- nies were discovered in close proximity to M. andrei nests. In one instance at Hop- land Research and Extension Center, a mature colony of P. creightoni was located Volume 14, Number 2, 2005 149 □ D P. creishtoni ♦ P. californica □ [J D P. creightoni ♦ P. creightoni-likc variant of P. californica\ Figs 24-25. Known distributions in northern California, western Nevada, and southern Oregon of P. nica and P. creightoni (24), and P. creightoni and the P. creightoni-hke variant of P. californica (25). califor- only 20cm from the entrance of a large and active colony of M. andrei, the Pheidole nest being surrounded by the chaff piles and foraging trails of M. andrei. Colonies of P. californica were never observed in close proximity to nests of M. andrei. Giv- en that P. creightoni and M. andrei are both seed harvesting species, and thus potential competitors, it seems paradoxical that they would preferentially nest in close proximity to one another. Pheidole clementensis. — This species is found in arid and semi-arid habitats of southern California and northern Baja Cal- ifornia (Fig. 23) at moderate elevations (160 to 800m, mean = 530m). Based on collection data from specimens examined, nesting is most often beneath stones, but the author has not made any personal ob- servations. DISCUSSION AND CONCLUSIONS Pheidole californica is the most variable member of the californica complex at both the local and geographic scale, with marked differences in morphology among populations, among nests within a popu- lation, and among the members of a single colony. In spite of this variability, which has led to poor characterization of the spe- cies and confusion with other members of the complex, I have identified some traits that consistently separate P. californica from its closest relatives. High levels of morphological variation in P. californica may be correlated with the large range of the species. It is possible that barriers to dispersal within the wide geographical range of P. californica have led to divergence in this species, while the comparatively small range of P. creightoni 150 Journal of Hymenoptera Research and P. clementensis has led to a corre- spondingly lower amount of variation. Another possible source of variation in the California! complex is hybridization be- tween species. At several localities P. calx- fornica and P. crcigJitoni are sympatric, and at one of these sites there is some evidence of hybridization. Thus, it may be that the confusing variation among and within some populations of P. California! is partly due to the influence of introgression with P. creightoni. Genetic work may be re- quired for the resolution of this and other questions related to variation in members of the californica complex. ACKNOWLEDGEMENTS I would like to thank Philip Ward (PSWC and UCDC) and Roy Snelling (LACM) for providing ac- cess to mateinal in collections. Useful material was also provided by Alex Wild and Emma Underwood. I want to specially thank my advisor Philip Ward for inspiration and guidance throughout this project. The following organizations helped make this project pos- sible by granting permission to collect ants on their land, and by providing additional assistance with field logistics: the Pyramid Lake Paiute Tribe, The Nature Conservancy (Dye Creek Preserve, Lassen Foothills), and The University of California (Hopland Research and Extension Center). Funding for the pro- ject was provided by the President's Undergraduate Fellowship program at the University of California, Davis. LITERATURE CITED Bolton, B. 1995. A new general catalogue of the ants of the world. Harvard University Press, Cambridge, Mass. 504 pp. Cole, A. C. 1933. Descriptions of two new ants of the genus Pheidole (Hymenoptera: Formicidae). An- nals of the Entomological Society of America 26: 616- 618. Cole, A. C. 1936. An annotated list of the ants of Ida- ho (Hymenoptera: Formicidae). The Canadian En- tomologist 68: 34-39. Creighton, W. S. 1950. The ants of North America. Bulletin of the Museum of Comparative Zoology at Harvard College 104: 172-174. Emery, C. 1895. Beitrage zur Kenntniss der norda- merikanischen Ameisenfauna. Zoologische jahr- biichcr. Abthcilung fiir Systematik, Geographic und Biologie der Tiere 8: 257-360. Emery, C. 1922. Hymenoptera: Fam. Formicidae, su- bfam. Myrmicinae. Pp. 95-206 in: Wytsman, P. Genera insectorum. Fasc. 174B. Bruxelles. Forel, A. 1901. Varietes myrmecologiques. Annals de la societe Entomologique de Bclgique 45: 334-382. Gregg, R. E. 1955. A new species of ant belonging to the Pheidole pilifera complex (Hymenoptera: For- micidae). Psyche 62: 19-28. Gregg, R. E. 1959. Key to the species of Pheidole (Hy- menoptera: Formicidae) in the United States. journal of the New York Entomological Society 66: 7-48. Gregg, R. E. 1969. New species of Pheidole from the Pacific coast islands (Hymenoptera: Formicidae). Entomological News 80: 93-101. Mayr, G. 1870. Neue Formiciden. Verhandlungeu der k.k. Zoologisch-Botanischen Gescllschaft in Wein 20: 939-996. Ward, P. S. 1999. Deceptive similarity in army ants of the genus Neivatm/rmex (Hymenoptera: For- micidae): taxonomy, distribution and biology of N. californicus (Mayr) and N. nigrescens (Cresson). Journal of Hymenoptera Research 8: 74-97. Ward, P. S. 2000. On the identity of Pheidole vaslitii Pergande (Hymenoptera: Formicidae), a neglect- ed ant from Baja California. Journal of Hymenop- tera Research 9: 85-98. Wheeler, G. C. and Wheeler, J. N. 1972. Ant larvae of the subfamily Myrmicinae. Pan-Pacific Entomolo- gist 47: 245-256. Wheeler, G. C. and Wheeler, J. N. 1986. The ants of Nevada. Natural Histon/ Museum of Los Angeles County. Wheeler, W. M. 1915. Some additions to the North American ant-fauna. Bulletin of the American Mu- seum of Natural History 34: 389-421. Wilson, E. O. 2003. Pheidole in the New World, a dom- inant, hyperdiverse ant genus. Harvard University Press, Cambridge, Mass. 794 pp. J. HYM. RES. Vol. 14(2), 2005, pp. 151-176 A Review of the Species of the New World Braconid Genus Cyclaulacidea (Hymenoptera) with Key and Descriptions of Nine New Species Jason W. Leathers/ Darlene D. Judd, and Andrew V. Z. Brower Oregon State University, Department of Zoology, 3029 Cordley Hall, Corvallis OR 97331 USA "Author for correspondence: jleathers@cdfa.ca.gov Abstract. — The Neotropical parasitic wasp genus Cyclaulacidea Quicke & Delobel (Hymenoptera: Braconidae) contains two previously described species from Peru and Brazil that are known to feed on Bruchidae and Curculionidae associated with palms. An examination of Neotropical bra- conines from 19 insect collections reveals that species of Cyclaulacidea are much more widespread. Nine new species of Cyclaulacidea are described, and an identification key to the eleven known species is provided. The new species are: C. pottsae from Mexico; C. adairae, C. hunteri, and C. snyderorum from Costa Rica; C. fergusoni from Panama; C. picki, C. rominus, and C. sharkeyi from Suriname; and C. riceorum from Peru and Brazil. The range of C. bruchivorus Quicke is expanded from Peru to include Brazil, Bolivia, and Suriname, and that of C. matilei Villemant and Simbolotti is expanded from Brazil to Colombia. Cyclaulacidea Quicke and Delobel cur- contains 33 described species that share a rently contains two species of wasps that unique facial sculpture of paired ridges are ectoparasitic on Coleoptera feeding on that run from the antennal sockets to the palms (Quicke 1997, Villemant and Sim- clypeus (Figs. 2, 3, 4) and which are divid- bolotti 2000). Cyclaulacidea bruchivorus ed into seven genera: Compsobracon, Calob- Quicke has been reared in Peru from Car- racon Szepligeti, Cyclaulax Cameron, yoborus serripes Sturm (Bruchidae: Pachy- Compsobraconoides, Cyclaulacidea, Gracili- merinae) feeding on fruits fallen from As- bracou Quicke, and Sacirema Quicke trocaryum javarense Trail ex Drude, A. (Quicke 1997). chouta Martins, and A. macrocah/x Burret (Quicke and Delobel 1995). Cyclaulacidea METHODS matilei Villemant and Simbolotti has been Specimens examined. — As part of a ge- reared from Foveolus sp. (Curculionidae: neric-level revision of the entire Compso- Rhynchophorinae: Sphenophorini) feed- bracou group, 4,918 specimens of Neotrop- ing in floral bracts of Euterpe oleracea C. ical Braconinae were borrowed from 19 in- Martius (Villemant and Simbolotti 2000). sect collections. Additionally, the senior Neither species has been documented out- author recently had the opportunity to ex- side of its type locality country. amine type specimens of C. bruchivorus Cyclaulacidea is part of the Compsobracon and C. matilei during a visit to the Musee Ashmead group of Neotropical parasitic National d'Histoire Naturelle in Paris, wasps (Braconidae: Braconinae), a diverse Specimens from the Compsobracon group group of at least 100 species that vary used in this study are marked with unique from the enormous, brightly colored spe- numbers on their determination labels in cies of Compsobracon to the tiny and incon- the format JL###### to allow the contin- spicuous species of Compsobraconoides ued association of notes and observed Quicke. The Compsobracon group currently character states with particular specimens. 152 Journal of Hymenoptera Research Table 1. Summary of meristic and continuous measurements in species of Cyclaulacidea. L width, D = distance, H = height, FW = Fore wing. length, W adai. brttc. ferg. hunt. Length (mm) 7.0-8.3 7.0-12.7 8.6 7.0-7.5 Flagellomeres 42-45 49-63 46 42 Scape L:W 1.5-2.0 1.7-2.2 2.1 1.8-2.1 First:second flagellomere 1.2-1.4 1.2-1.7 1.4 1.3-1.4 Firstrthird flagellomere 1.3-1.6 1.3-1.7 1.5 1.4-1.5 Third flagellomere L:W 1.0-1.3 0.9-1.3 1.5 1.1 Apical flagellomere L:W 1.5-1.8 1.8-2.2 1.8 1.6 Horizontal L eye: L of head behind eye 1.6-2.2 0.9-1.2 1.6 1.5 Diameter posterior ocellus: post-ocellar L 0.9-1.5 1.0-1.5 0.8 1.0-1.2 D between posterior ocellus and eye: post-ocellar L 2.9-3.8 3.4-4.3 2.6 3.9-4.9 Eye H:W 1.2-1.3 1.3-1.4 1.2 1.3-1.4 Eye H:W of face 1.2-1.5 1.1-1.6 1.5 1.4 W of head:W of face 2.2-2.8 2.0-2.8 2.6 2.4 Inter-tentorial D:clypeus H 2.0-2.4 1.8-2.2 2.4 2.2-2.3 Tentorio-ocular Dxlypeus H 0.8-1.1 0.8-1.2 0.9 1.1 Face W:H 0.9-1.2 1.0-1.2 0.9 0.9-1.0 Malar space:eye H 0.2-0.3 0.1-0.2 0.2 0.1-0.2 Mesosoma L:H 1.3-1.5 1.4-1.7 1.6 1.4 Propodeal spiracle H:W 2.5-3.3 2.0-2.7 2.3 2.0-2.2 Foretibia L:forefemur L 1.1-1.2 1.1-1.2 1.0 1.1 Foretarsus L:forefemur L 1.5-1.6 1.5-1.7 1.6 1.6 Fore basitarsus L:W 3.7-5.0 3.9-5.7 4.7 4.2-4.8 Fore basitarsus L:second tarsomere L 1.6-1.8 1.5-1.8 2.0 1.6-1.7 Hind femur L:W 3.8-1.2 3.7-4.0 4.0 3.6-4.0 Hind femur L:hind basitarsus L 2.0-2.4 1.6-1.9 1.9 1.8-2.7 Hind tibia:hind basitarsus L 2.5-2.9 2.3-2.8 2.7 2.4-2.7 Outer tibial spur L:basitarsus L 0.5-0.6 0.3-0.5 0.4 0.5 Inner tibial spur L:basitarsus L 0.6-0.7 0.6 0.6 0.7 Hind basitarsus L:W 3.4-4.1 5.4-7.3 4.6 4.0-4.7 FW 1M:1RS L 1.7-2.5 1.5-2.3 2.4 1.9-2.0 FW 2M:r-m L 3.5-4.1 3.1-4.1 4.2 3.4-3.5 FW lm-cu:r L 1.2-1.6 1.0-1.4 1.3 1.1-1.2 FW lm-cu:(RS+M)a thickness 1.5-2.2 1.0-2.0 1.5 1.2-1.4 FW 2RS:r-m L 1.3-1.5 1.0-1.2 1.5 1.1-1.2 FW 3RSa:r-m L 3.0-3.4 2.7-3.6 3.3 3.0 FW 3RSa:r L 4.6-6.2 4.5-5.7 4.8 4.8-5.3 FW 3RSb:r L 6.0-7.6 4.4-6.0 5.8 5.1-5.5 FW L 6.9-8.0 8.6-12.0 7.8 7.0-7.5 Hind wing vein Rla:lr-m L 1.3-1.8 1.1-1.4 1.4 1.5-1.6 First tergite L:W 0.9-1.1 0.7-1.2 0.8 0.8-0.9 Second tergite L:W 0.4-0.5 0.4-0.5 0.5 0.4-0.5 Second tergite L:third tergite L 0.7-0.9 0.6-0.8 0.7 0.8-0.9 Third tergite L:W 0.5-0.6 0.5-0.7 0.7 0.5-0.6 Ovipositor:body L 0.8-1.0 1.4-1.5 6 1.1-1.2 These numbers are included for type spec- imens in the species descriptions and are included in the distribution section (Ap- pendix 1) for new specimens of C. bruchi- vorus and C. matilei. Because resources for the identification of New World braconines are sparse and Cyclaulacidea was only recently described, specimens of Cyclaulacidea are rare in col- lections. Several of the species described herein are represented only by single specimens. Single specimens were recog- Volume 14, Number 2, 2005 153 Table 1. Continued. matt pick. yott. rice. rami. shut >n\id. 7.0-9.5 10.3 8.0 7.8-10.1 8.8-9.6 8.8-14.1 7.2-9.2 46-54 BR 46 48-52 48-53 54-58 46-52 1.3-2.5 1.8 1.5 1.7-2.1 1.6-1.7 1.9-2.3 1.8-1.9 1.3-1.4 1.3 1.4 1.3-1.5 1.2-1.4 1.2-1.5 1.3-1.4 1.4-1.5 1.3 1.5 1.4-1.6 1.3-1.4 1.4-1.6 1.4 1.1 1.5 1.1 1.2-1.3 1.0-1.1 1.1-1.3 1.1-1.3 1.6-1.8 BR 1.7 1.8-2.9 1.8-1.9 1.8-2.3 1.8-1.9 1.7 0.9 1.5 1.0-1.4 1.4-1.7 0.7-0.8 1.7 1.2 1.6 0.8 0.9-1.2 1.0-1.1 0.9-1.4 0.9-1.1 4.3 5.3 2.9 2.8-4.0 3.5 3.8-5.6 2.9-3.3 1.2 1.3 1.3 1.3-1.4 1.3 1.3-1.5 1.3 1.1-1.3 1.3 1.2 1.2-1.7 1.2 1.0-1.2 1.3 2.0-2.3 2.4 2.3 2.1-2.7 2.2 1.9-2.3 2.3-2.4 2.7-3.2 2.5 2.5 1.9-3.1 2.0-2.3 1.7-2.3 2.5-2.8 1.0-1.5 1.1 1.3 0.7-1.1 1.0-1.1 1.0-1.3 1.1-1.2 1.3 1.0 1.1 0.7-1.1 1.1-1.2 1.1-1.2 1.0-1.1 0.2 0.2 0.2 0.1-0.2 0.2 0.2-0.3 0.2 1.5-1.9 1.6 1.5 1.6-1.7 1.4-1.5 1.4-1.7 1.5-1.6 3.3 1.9 2.0 2.0-2.5 2.7-3.2 1.8-2.4 1.9-2.3 1.1-1.2 1.1 1.2 1.1-1.2 1.1-1.2 1.0-1.2 1.1-1.2 1.4-1.6 1.7 1.6 1.6-1.7 1.6-1.7 1.5-1.7 1.6-1.7 4.3 5.2 4.2 4.3-5.4 3.3-4.3 3.9-5.7 4.5-5.1 1.7 1.7 1.8 1.5-1.8 1.6-1.7 1.7-1.9 1.5-1.8 4.0 4.1 3.8 3.8-4.0 3.7-4.1 3.8-4.2 4.0-4.4 1.8 1.5 1.8 1.6-1.8 1.9-2.1 1.6-1.8 1.7-1.8 2.4 2.6 2.6 2.3-2.7 2.5-2.6 2.5-2.6 2.4-2.6 0.4-0.5 0.3 0.4 0.4-0.5 0.5 0.4 0.4-0.5 0.6 0.5 0.6 0.6 0.6-0.7 0.5-0.6 0.6 6.0 5.5 5.5 5.0-6.2 4.2-1.5 5.2-6.2 4.4-5.2 1.7 2.3 2.1 1.9-2.1 1.5-1.7 1.8-2.1 1.7-2.1 3.3 3.5 3.6 3.3-3.8 3.3-3.8 3.1-3.6 3.2-3.5 1.1-1.3 1.3 1.1 1.0-1.5 1.0 1.2-1.6 0.9-1.3 1.6 1.7 1.8 1.1-2.2 1.5-2.0 1.1-1.7 1.5-1.6 1.1 1.2 1.1 1.2-1.3 1.0-1.2 1.0-1.2 1.1-1.2 2.9 2.9 3.2 2.7-3.2 2.8-3.1 2.9-3.2 2.8-2.9 4.1-4.7 4.8 5.1 4.3-6.2 4.0-4.4 5.0-6.3 3.8-5.4 4.9-5.7 5.4 4.7 5.2-7.2 4.6-5.0 4.6-6.0 4.1-5.4 9.7 9.0 7.6 7.3-8.6 9.0-10.3 8.7-12.8 7.0-9.1 1.5-1.8 1.8 1.5 1.4-1.6 1.4 1.3-1.5 1.2-1.4 0.9-1.2 1.1 0.9 0.8-1.1 0.9-1.0 1.1-1.3 0.8-0.9 0.4-0.8 0.7 0.5 0.5-0.6 0.4-0.5 0.5-0.6 0.4-0.5 0.7 1.4 0.7 0.7-0.9 0.7-0.9 0.7-0.9 0.7-0.8 0.4 0.5 0.6 0.6-0.7 0.5-0.6 0.5-0.7 0.5-0.6 1.0 1.1 1.0 1.2 0.9 0.8-1.1 1.1 nized as new species only when they pos- sessed unique combinations of at least several characters and had a disjunct geo- graphic distribution. All of the known specimens of Cydaidacidcn are in good condition, although a few of the older ones are moderately dusty (e.g., Fig. 3B). Several museums have large collections of unsorted Neotropical braconines (e.g., the American Entomological Institute). There are undoubtedly additional species of Cyclaulacidea that await description RESULTS 154 Journal of Hymenoptera Research when funding is available to curate these (CAS); Entomological Museum, Utah State accessions. University, Logan, Utah, USA (EMUS); Rocky Mountain Systematic Entomology Morphology. — Morphological terminolo- Laboratory, University of Wyoming, Lar- gy and character systems examined follow amie, Wyoming, USA (ESUW); Institute Sharkey and Wharton (1997), with the ex- Alexander von Humboldt, Santafe de Bo- ception of morphometric characters, gota, Colombia (IAVH); Musee National which follow van Achterberg (1979). As d'Histoire Naturelle, Paris, France part of a larger study of the Compsobracon (MNHN); Museu de Zoologia da Univer- group of New World Braconinae, all spec- sidade de Sao Paulo, Sao Paulo, Brazil imens of Cyclaulacidea were examined for (MZSP); Oregon State Arthropod Collec- a total of 68 discrete, 44 continuous, and 3 tion, Oregon State University, Corvallis, meristic morphological characters, as well Oregon, USA (OSUO); Nationaal Natu- as 41 color characters. Continuous char- urhistorische Museum, Leiden, Nether- acters were measured using a Microcode lands (RMNH); Department of Entomolo- II (Boeckeler Instruments). Length was gy Insect Collection, Texas A&M Univer- measured from head to abdomen. Mea- sity, College Station, Texas, USA (TAMU); sures of face height were measured from National Museum of Natural History, the tentorial pits rather than the top of the Smithsonian Institution, Washington D.C., clypeus. Meristic and continuous data is USA (USNM). included in the species descriptions and summarized in Table 1. The angle H of Fore wing veins C+SC + R and IRS was Of the 4,918 specimens of Braconinae estimated using the formula tan-'e = examined, 1,133 are members of the Comp- ((distance from intersection of IRS and sobracon group, but only 41 are members (RS + M)a to C+SC + R in a basad direction of Cyclaulacidea. Additional specimens of measured perpendicular to IRS) / (length C. bruchivorus and C. matilei are among of IRS)) (Fig. 6A). The species descriptions these specimens, as are 9 species new to include character states from both holo- science. No information on host associa- types and paratypes. When there is vari- tion or other ecological data are recorded ation in this data holotype information is on tne labels of any specimens of the new denoted in square brackets. species. In many braconines the clypeus is sep- DISTRIBUTION arated from the rest of the face by a raised ridge (as in Fig. 4A). Furthermore, in some Specimens of Cyclaulacidea, previously species of Cyclaulacidea the clypeus is also documented from only Peru and Brazil, elevated such that parts of it are level with are reported from Mexico, Costa Rica, this ridge. In the species descriptions Panama/ Colombia, Suriname, and Bolivia 'clypeus partially filled in dorsally' means (Fi8- *)■ The ran§e of C- bruchivorus is ex- that the part of the clypeus closest to the Panded from Peru to include Brazil, Boliv- face is level with this ridge and the part ia' and Surinarrie (Fig. 1, Appendix 1). The closest to the labrum appears excavated only new specimen of C. matilei bears col- and is not level with the ridge (as in Fig lection information from Colombia (Fig. 1, 2B). Appendix 1). Depositories.— Specimens of Cyclaulaci- SYSTEMATICS dea were found in the following museums The acronyms used here are taken from Cyclaulacidea Quicke & Delobel Arnett et. al. (1993): California Academy Cyclaulacidea Quicke & Delobel, 1995: 218-219. of Sciences, San Francisco, California, USA Diagnosis.— Species of Cyclaulacidea can Volume 14, Number 2, 2005 155 Legend ♦ adairae 0 bruchivorus * fergusom T hunteri A matilei + picki e pottsae * riceorum ft rominus <•> sharkeyi X snyderorum N W ^^ E S Fig. 1. Distribution map for species of Cyclaulacidea. be distinguished from other genera of Bra- coninae using the key of Quicke (1997) and by the presence of the following pu- tative synapomorphies: a median ridge on the face is developed into a raised tear- drop, chevron, or butterfly-shaped area (Figs. 2, 3, 4) and the first tergite has a strongly raised median rectangular to tongue-shaped area (Figs. 8A, 8B, 8C). One of the new species, C. adairae, keys to Compsobraconoid.es in Quicke's key due to the shape of its scape; however, this taxon is much larger than any species of Comp- sobraconoides and possesses the tear-drop shaped area on the face and rectangular bump on the first tergite characteristic of Cyclaulacidea. All known species of Cyclaulacidea also possess the following combination of char- acters: HEAD: Pedicel not swollen or heavily sclerotized. Scape lacking basal concavity. First flagellomere swollen basally toward vertex of head. Apical flagellomere acicu- late. Antennal sockets not extended from head, lacking plate-like shelf and enlarged sockets. Vertex of head with smooth de- pression and groove medially. The face is glabrous and has a pair of main, subme- dial, longitudinal ridges running from the clypeus to the antennal sockets (Figs. 2, 3, 4). Area between ridges and eyes with lad- der-like series of horizontal carinae. Clyp- eus separated from rest of face by rugose ridge. MESOSOMA: Metanotum mostly smooth. Propodeum lacking median lon- gitudinal carina. Tarsal claw with very small basal lobe. Hind tibia lacking lon- gitudinal depression. Hind telotarsus lack- ing especially thick setae. Fore wing vein (RS+M)a is strongly curved (Fig. 6). Hind wing with 1 basal hamulus and an area of reduced setosity apicad to vein cu-a. 256 Journal of Hymenoptera Research METASOMA: First tergite trapezoidal, branch of suturiform articulation absent, lacking median longitudinal and Y- Third tergite smooth, lacking pinched-up shaped carinae. Second median tergite area, median longitudinal carina, and lacking raised mid-basal triangular area mid-basal triangular area. Hypopygium pointing posteriorly or anteriorly. Apical pointed apically. KEY TO SPECIES OF CYCLAULACIDEA 1. Fore wing entirely black (as in Fig. 10B) or black with one clear band (Fig. 12B) 2 Fore wing with two yellow (as in Fig. 11) or clear bands (as in Fig. 12C) 5 2(1). Fore wing with one clear band in apical third; maxillary and labial palpomeres black basally, yellowish orange apically; (Fig. 12B) C. pottsac n. sp. Fore wing entirely black (as in Fig. 10B); maxillary and labial palpomeres entirely black or entirely white 3 3(2). Terga 4-6 black dorsally, yellowish orange laterally; maxillary and labial palpomeres entirely white; suturiform articulation represented by a deep groove (Fig. 7A); fore tarsus strongly laterally compressed (Figs. 5A, 5B); (Fig. 10B) C. fergusoni n. sp. Terga 4-6 entirely black; maxillary and labial palpomeres entirely black; suturiform articulation represented by weak groove (as in Fig. 7B); fore tarsus not strongly laterally compressed (as in Figs. 5C, 5D) 4 4(3). Scape longer dorsally than ventrally (Fig. 5E); rectangular bump on petiole narrowing posteriorly (Fig. 8A); main, submedial, longitudinal facial ridges bowed outward (Fig. 2A); median carina on face developed into tear-drop shaped area medially (Fig. 2A); (Fig. 10D) C. adairae n. sp. Scape longer ventrally than dorsally (Fig. 5F); rectangular bump on petiole not narrow- ing posteriorly (Fig. 8B); main, submedial, longitudinal facial ridges diverging out- ward straight from clypeus to antennal sockets (Fig. 3A); median carina on face developed into chevron to butterfly-shaped area medially (Fig. 3A); (Fig. IOC) . . . C. snyderorum n. sp. 5(1). Mesosoma entirely yellowish orange 6 Mesosoma entirely black, or mostly black with some yellowish orange on dorsal surface and around margins of pronotum, tegula, and /or sternaulus 7 6(5). Suturiform articulation barely distinguished from terga 2 and 3, lacking groove (Fig. 7C); bump on petiole tongue-shaped, wider posteriorly than anteriorly (Fig. 8C); main, submedial, longitudinal facial ridges bowed outward (as in Figs. 2A, 2B); ovipositor sheath entirely black; (Fig. 12C) C. picki n. sp. Suturiform articulation with shallow groove (as in Fig. 7B); bump on petiole rectan- gular (as in Fig. 8B); main, submedial, longitudinal facial ridges parallel (Fig. 3B) or diverging straight from clypeus to antennal sockets (as in Fig. 3A); ovipositor sheath black with some yellowish orange in apical third (but black at apical tip); (Fig. 12A) C. sharkeyi n.sp. 7(5). Forecoxa usually entirely yellowish orange, sometimes with some black basally; mid femur entirely yellowish orange; Fore wing vein lcu-a intersects Cu distad 1M (as in Fig. 6A) g Forecoxa usually entirely black, sometimes yellowish orange; mid femur usually en- tirely black, sometimes with yellowish orange on basal and apical ends, or entirely yellowish orange; Fore wing veins 1M and lcu-a intersect (interstitial) (as in Fig. 6B) 9 Volume 14, Number 2, 2005 157 8(7). Inter-tentorial distance 1.8-2.2 times greater than clypeus height (Fig. 4A); maxillary and labial palpomeres entirely yellowish orange to white; (Fig. 11 A) . . C. bruchivorus Inter-tentorial distance 2.7-3.2 times greater than clypeus height (Fig. 4B); maxillary and labial palpomeres black basally, yellowish orange apically; (Fig. 11C) . . C. matilei 9(7). Costa yellow; fore tibia entirely yellowish orange; (Fig. 11B) C. riceorum n. sp. Costa black; fore tibia entirely black, or mostly black with some yellowish orange in basal and/or apical sixths 10 10(9). Facial ridges parallel (as in Fig. 3B); second tergite with slightly elevated pinched-up area anteriorly (Fig. 9A); antenna with less than 46 flagellomeres; (Fig. 10 A) C. hunteri n. sp. Facial ridges bowed outward (Fig. 2B); second tergite with strongly pinched-up area anteriorly (Fig. 9B); antenna with 48 to 53 flagellomeres; (Fig. 11D) C. romimis n. sp. SPECIES DESCRIPTIONS Cyclanlacidea adairae Leathers n. sp. Figs. 2A, 5C, 5D, 5E, 7B, 8A, 10D Diagnosis. — Scape longer dorsally than ventrally (Fig. 5E). Rectangular bump on petiole narrowing posteriorly (Fig. 8A). Terga 1-3 black dorsally, yellowish orange laterally (Fig. 10D). Length.— 7.0-8.3 [8.0] mm. Head. — Antenna with 42-45 [44] flagel- lomeres. Scape longer dorsally than ven- trally. Scape lacking apical and pre-apical shelf-like process, [1.5]-2.0 times longer than maximally wide. First flagellomere 1.2-1.4 [1.3] times longer than second fla- gellomere, 1.3— [1.6] times longer than third flagellomere. Third flagellomere 1.0- 1.3 [1.1] times longer than wide. Apical flagellomere 1.5-1.8 [1.6] times longer than wide. Flagellomere length equal to or greater than width. Horizontal length of eye 1.6-[2.2] times longer than length of head behind eye. Transverse diameter of posterior ocellus 0.9-1.5 [1.1] times post- ocellar length. Distance between posterior ocellus and eye 2.9-3.8 [3.5] times post- ocellar length. Longitudinal bump be- tween antennal sockets present. Facial ridges bowed outward. Anterior groove between antennal sockets absent. Area be- tween ridges filled in creating a raised me- dian area. Median carina on face present, developed into raised tear-drop shaped Fig. 2. Face of A) C. adairae (JL000100) and B) C. rominus (JL000228). Both have facial ridges that are bowed outward and a raised tear-drop shaped area in the center of the face. 158 Journal of Hymenoptera Research >^mFI ► ^^^^ 1 wm 1 •J£«8i|E\ 1 ill T; f 'Ml r/ w.^ B * Fig. 3. Face of A) C. snyderorum (JL000101) with fa- cial ridges that are diverging outward from the clyp- eus to the antennal sockets and B) C. sharkeyi (JL000203) with facial ridges that are more parallel. Both have raised chevron to butterfly-shaped areas in the center of the face. area. Area between median carina and ridges with ladder-like series of horizontal carinae. Ridges running at 45° angle from middle ridge to antennal sockets strong. Groove around eyes present, [smooth] or crenulate. Eye height 1.2— [1.3] times great- er than eye width, [1.2]-1.5 times greater than width of face. Width of head 2.2-2.8 [2.3] times greater than width of face. In- ter-tentorial distance 2.0-2.4 [2.2] times clypeus height. Tentorio-ocular distance 0.8-[l.l] times clypeus height. Clypeus Fig. 4. Face of A) C. bruchivorus (JL000047) and B) C. matilei (JL000201). The arrow points to the clypeus, the shape of which can be used to distinguish be- tween the two species. CH = clypeus height, ITD = inter-tentorial distance. [partially filled in dorsally], or completely filled in but uneven. Ventral margin of clypeus concave. Area around clypeus with series of large crenulae and sharp ridge separating from rest of face, or [sep- arated from rest of face by large smooth groove]. Face 0.9-1.2 [1.0] times wider than high. Malar suture paralleled by 2- [5] ridges. Malar space [0.2]-0.3 times eye height. Mesosoma. — Mesosoma 1.3-1.5 [1.4] times longer than high. Pronotum with deep, smooth groove anteriorly. Notaulus not impressed. Border between mesoscu- tum and scutellum with weak carinate Volume 14, Number 2, 2005 159 Fig. 5. A) Dorsal and B) lateral images of fore tarsus of Cfergusoni (JL000235) and C) dorsal and D) lateral images C. adairae (JL000102); arrow indicates relative lateral compression in fore tarsus of C. fergusoni. Scape of E) C. adairae (JL000102) and F) C. snyderorum (JL000107); arrow denotes ventral surface which is not longer than dorsal surface in C. adairae but is lon- ger than dorsal surface in C. snyderorum. groove, lacking strong median carina and enlarged median pit. Propodeal spiracle oval to [crescent-shaped], 2.5-3.3 [2.9] times higher than wide. Fore tibia [1.1]— 1.2 times longer than fore femur. Fore tarsus not laterally com- pressed, [1.5]— 1.6 times longer than fore femur. Fore basitarsus [3.7]-5.0 times lon- ger than wide, 1.6— [1.8] times longer than second tarsomere. Hind femur 3.8-4.2 [3.9] times longer than wide, 2.0-[2.4] times longer than basitarsus. Hind tibia 2.5-[2.9] times longer than basitarsus. Outer and inner hind tibial spurs 0.5-[0.6] and 0.6-[0.7] times longer than basitarsus, respectively. Hind basitarsus 3.4-4.1 [3.6] times longer than wide. Fore wing venation: 1M and lcu-a in- tersect. (RS + M)b broken apically. 1M 1.7- 2.5 [2.0] times longer than IRS. 2M 3.5-4.1 [3.6] times longer than r-m. lm-cu 1.2-1.6 [1.4] times longer than r, 1.5-2.2 [1.6] times as thick as (RS+M)a. 2RS [1.3J-1.5 times longer than r-m. 3RSa [3.0]-3.4 times lon- ger than r-m, 4.6-6.2 [5.3] times longer than r. 3RSb 6.0-7.6 [7.1] times longer than r. C + SC + R and IRS forming an angle of 67-73° [71°]. Fore wing length 6.9-8.0 [7.9] mm. Hind wing vein Rla [1.3]— 1.8 times lon- ger than lr-m. Metasoma. — Base of petiole deeply ex- cavated. First tergite with strongly raised rectangular bump. First tergite with lateral carina closely paralleling median bump but not forming notches, with pair of lat- eral carinae entirely separated from me- dian bump. First tergite [0.9J-1.1 times longer than wide. Border between first and second tergite straight with edges curving anteriorly. Second median tergite with strongly pinched-up area not reach- ing third tergite. Suturiform articulation with weak, smooth, M-shaped, weakly arched groove; with carina present along anterior margin. Second tergite [0.4]-0.5 times longer than wide, [0.7]-0.9 times longer than third tergite. Third tergite [0.5]-0.6 times longer than wide. Hypo- pygium with convex dorsal margin. Ovi- positor 0.8-[1.0] times body length. Color. — Entirely black except lateral parts of terga 1-3 yellowish orange and sometimes apical parts of mid and hind trochantellus yellowish orange or red. Wings entirely black. Distribution and material examined. — Known from Guanacaste, Alajuela, and Heredia provinces of Costa Rica. 79 9 16 examined. Holotype 9. COSTA RICA: Guanacaste: 3km SE R. Naranjo, 3-8.iii.1992, F.D. Parker (EMUS— JL000100). Paratypes. COSTA RICA: Guanacaste: 3km SE R. Naranjo: 29$, 3-8.iii.1992, F.D. Parker (EMUS— JL000102, 000105); 19, 28.xi-5.xii.1991, F.D. Parker (EMUS— JL000103); 16, xii.1991, F.D. Parker (EMUS— JL000106). Alajuela: 29 9, 20.xi.1990, F.D. Parker (EMUS— JL000231, 000232). Heredia: F. La Sel- va, 3km S. Pto. Viejo: 19, 14.iii.1980, H.A. Hespen- heide (ESUW— JL000233). 160 Journal of Hymenoptera Research Fig. 6. Fore wing of A) C. bruchivorus (JL000246) and B) C. riceorum (JL000162). 6 = the angle of Fore wing veins C+SC+R and IRS. Remarks. — Coloration appears almost identical to C. snyderorum sp. n., but can be consistently distinguished using the di- agnostic characters in the key. Etymology. — For Lila Adair for her sup- port of science education at Central Gwin- nett High School. Cyclaalacidea fergusoni Leathers n. sp. Figs. 5A, 5B, 7 A, 10B Diagnosis. — Suturiform articulation with deep groove (Fig. 7A). Facial ridges bowed inward. Terga 4-6 black dorsally, yellowish orange laterally (Fig. 10B). Length. — 8.6 mm. Head. — Antenna with 46 flagellomeres. Scape longer ventrally than dorsally. Scape with shelf-like process apically, lacking pre-apical shelf, 2.1 times longer than maximally wide. First flagellomere 1.4 times longer than second flagello- mere, 1.5 times longer than third flagel- lomere. Third flagellomere 1.5 times lon- ger than wide. Apical flagellomere 1.8 times longer than wide. Flagellomere length equal to or greater than width. Horizontal length of eye 1.6 times longer than length of head behind eye. Trans- verse diameter of posterior ocellus 0.8 times post-ocellar length. Distance be- tween posterior ocellus and eye 2.6 times post-ocellar length. Longitudinal bump between antennal sockets absent. Facial ridges bowed inward. Deep anterior groove between antennal sockets absent. Area between ridges filled in creating a raised median area. Median carina on face present, developed into raised tear- drop shaped area. Area between median carina and ridges with ladder-like series of horizontal carinae. Ridges running at 45° angle from middle ridge to antennal sockets strong. Groove around eyes pre- sent and smooth. Height of eye 1.2 times greater than eye width and 1.5 times greater than width of face. Width of head 2.6 times greater than width of face. Inter- tentorial distance 2.4 times clypeus height. Tentorio-ocular distance 0.9 times longer than clypeus high. Clypeus com- Volume 14, Number 2, 2005 161 Fig. 7. Metasomal syntergite 2 + 3 with A) deep groove on C. fergusoni (JL000235), B) shallow groove on C. adairae (JL000102), C) lack of groove on C. picki (JL000202). Arrow indicates suturiform articulation. pletely filled in but uneven. Ventral mar- ture paralleled by 6 ridges. Malar space gin of clypeus concave. Area around 0.2 times eye height. clypeus with series of large crenulae and Mesosoma.- — Mesosoma 1.6 times longer sharp ridge separating from rest of face, than high. Pronotum with deep, smooth Face 0.9 times wider than high. Malar su- groove anteriorly. Notaulus weakly im- ib2 Journal of Hymenoptera Research Fig. 8. First tergite of A) C. adairae (JL000232) with rectangular bump narrowing posteriorly, B) C. snyderorum (JL000107) with rectangular bump, and C) C. jridd (JL000202) with toungue-shaped bump. pressed. Border between mesoscutum and scutellum with carinate groove, lacking median carina and enlarged median pit. Propodeal spiracle crescent-shaped, 2.3 times higher than wide. Fore tibia 1.0 times the length of fore femur. Fore tarsus strongly laterally com- pressed, 1.6 times longer than fore femur. Fore basitarsus 4.7 times longer than wide, 2.0 times longer than second tarsomere. Hind femur 4.0 times longer than wide, 1.9 times longer than basitarsus. Hind tib- ia 2.7 times longer than basitarsus. Outer and inner hind tibial spurs 0.4 and 0.6 times longer than basitarsus, respectively. Hind basitarsus 4.6 times longer than wide. Fore wing venation: 1M and lcu-a in- tersect. (RS+M)b broken apically. 1M 2.4 times longer than IRS. 2M 4.2 times longer then r-m. lm-cu 1.3 times longer than r, 1.5 times as thick as (RS + M)a. 2RS 1.5 times longer than r-m. 3RSa 3.3 times lon- ger than r-m, 4.8 times longer than r. 3RSb 5.8 times longer than r. C + SC + R and IRS forming an angle of 67°. Fore wing length 7.8 mm. Hind wing vein Rla 1.4 times longer than lr-m. Metasoma. — Base of petiole deeply ex- cavated. First tergite with strongly raised rectangular bump. First tergite with lateral carina closely paralleling median bump but not forming notches, with pair of lat- eral carina entirely separated from median bump. First tergite 0.8 times longer than wide. Border between first and second ter- gite straight with edges curving anterior- ly. Second tergite with strongly pinched- up area not reaching third tergite. Suturi- form articulation with deep, smooth, M- shaped, weakly arched groove; carina along anterior margin present. Second ter- gite 0.5 times longer than wide, 0.7 times longer than third tergite. Third tergite 0.7 times longer than wide. Color. — Black except as follows: mouth- parts white, pronotum with yellowish or- ange stripe on ventral third, fore tarsus and mid tarsus yellowish orange with tel- otarsus black, hind trochantellus mostly black with some red apically, tergum 1 yellowish orange laterally, terga 2 and 3 reddish orange, terga 4-6 yellowish or- ange laterally. Wings entirely black. Distribution and material examined. — Known only from type specimen. 16* ex- amined. Holotype 3. PANAMA: N.Panama, 1050m, For- tuna, Chiriqui, 22-28.V.1979, H. Wolda (RMNH- JL000235). Etymology. — For George Ferguson for his generous support of this project and systematic entomology at Oregon State University. Volume 14, Number 2, 2005 163 Fig. 9. Second tergite of A) C. hunteri (JL000110) and B) C. rominus (JL000228). Arrow denotes pinched-up areas. Cyclaulacidea hunteri Leathers n. sp. Figs. 9A, 10A Diagnosis. — Fore wing banded: clear, black, clear, black (Fig. 10A). Maxillary and labial palpomeres white. Horizontal length of eye (in dorsal view) 1.5 times longer than length of head behind eye. Antenna with 42 flagellomeres. Length.— [7.0]-7.5 mm. Head. — Antenna with 42 flagellomeres. Scape longer ventrally than dorsally. Scape with shelf-like process apically, lacking pre-apical shelf. Scape [1.8J-2.1 times longer than maximally wide. First flagellomere [1.3]— 1.4 times longer than second flagellomere, [1.4]-1.5 times longer than third flagellomere. Third flagello- mere 1.1 times longer than wide. Apical flagellomere [1.6] times longer than wide. Flagellomere length equal to or greater than width. Horizontal length of eye 1.5 times longer than length of head behind eye. Transverse diameter of posterior ocel- lus 1.0-[1.2] times post-ocellar length. Dis- tance between posterior ocellus and eye 3.9-[4.9] times post-ocellar length. Facial ridges parallel. Anterior groove between antennal sockets very weak. Area between ridges filled in creating a raised median area. Median carina on face present, de- veloped into raised tear-drop shaped area. Area between median carina and ridges with ladder-like series of horizontal cari- nae. Ridges running at 45° angle from middle ridge to antennal sockets strong. Groove around eyes present and crenu- late. Eye height [1.3J-1.4 times greater than eye width, 1.4 times greater than width of face. Width of head 2.4 times greater than width of face. Inter-tentorial distance [2.2]-2.3 times clypeus height. Tentorio-ocular distance 1.1 times longer than clypeus high. Clypeus partially filled in dorsally. Ventral margin of clypeus concave. Area around clypeus with series of large crenulae and sharp ridge separat- ing from rest of face. Face [0.9]-1.0 times wider than high. Malar suture paralleled by 2 ridges. Malar space 0.1-[0.2] times eye height. Mesosoma. — Mesosoma 1.4 times longer than high. Pronotum with deep, smooth groove anteriorly. Notaulus not im- pressed. Border between mesoscutum and scutellum with carinate groove, lacking median carina or enlarged median pit; median part of groove sometimes filled in. 164 Journal of Hymenoptera Research Fig. 10. Automontage® lateral images of A) C. hunteri (JL000110), B) C.fergusoni (JL000235), C) C. snyderorum (JL000101), and D) C. adairae (JL000105). Ovipositor sheaths were removed from A. Magnification = 7.2X. Propodeal spiracle oval, 2.0-[2.2] times 1.6 times longer than fore femur. Fore bas- higher than wide. itarsus [4.2]-4.8 times longer than wide, Fore tibia 1.1 times longer than fore fe- [1.6]-1.7 times longer than second tarso- mur. Fore tarsus not laterally compressed, mere. Hind femur 3.6-[4.0] times longer Volume 14, Number 2, 2005 165 Fig. 11. Automontage® lateral images of A) C. bruchivorus (JL000246), B) C. riceorum (JL000162), C) C. matilei (JL000201), and D) C. rominus (JL000228). Ovipositor sheaths were removed from A, C, and D. Magnification = 4.7X. 166 Journal of Hymenoptera Research Fig. 12. Automontage* lateral images of A) C. sharkeyi (JL000236), B) C. pottsac (JL000243), and C) C. picki QL000202). Ovipositor sheaths were removed from B and C. Magnification = 5.1 X. Volume 14, Number 2, 2005 167 than wide, 1.8-[2. 7] times longer than bas- ajuela province of Costa Rica. 29 9 ex- itarsus. Hind tibia 2.4-[2.7] times longer amined. than basitarsus. Outer and inner hind tib- Holotype 9. COSTA RICA: Alajuela: 20km S. UPa- ial spurs .0.5 and 0.7 times length of basi- \a, 22-31. x. 1991, F.D. Parker (EMUS— JL000109). tarsus, respectively. Hind basitarsus 4.0- Paratype. 19, COSTA RICA: Alajuela: 20km S. Upala, [4.7] times longer than wide. 16*1990, F.D. Parker (EMUS-JL000110). Fore wing venation: 1M and lcu-a in- Etymology. — For Mark D. Hunter of The tersect. (RS + M)b broken apically. 1M 1.9- University of Georgia. [2.0] times longer than IRS. 2M 3.4-[3.5] times longer then r-m. lm-cu 1.1-[1.2] Cyclaulacideci picki Leathers n. sp. times longer than r, [1.2J-1.4 times as thick Fl8s- 7C' 8C' 12C as (RS + M)a. 2RS 1.1-[1.2] times longer Diagnosis. — Suturiform articulation than r-m. 3RSa 3.0 times longer than r-m, barely distinguished from remainder of 4.8-[5.3] times longer than r. 3RSb 5.1- metasomal syntergum 2 + 3 (Fig. 7C). [5.5] times longer than r. C + SC + R and Strongly raised bump on petiole tongue- 1RS forming an angle of 72°. Fore wing shaped (Fig. 8C). Median ridge on face de- length [7.0]-7.5 mm. veloped into tear-drop shaped area (as in Hind wing vein Rla [1.5]-1.6 times Ion- Figs 2A, 2B). ger than lr-m. Length. — 10.3 mm. Metasoma. — Base of petiole deeply ex- Head. — Antenna broken after 46th flagel- cavated. First tergite with strongly raised lomere. Scape longer ventrally than dor- rectangular bump. First tergite with lateral sally. Scape with shelf-like process apical- carina closely paralleling median bump ly, lacking pre-apical shelf, 1.8 times lon- but not forming notches, lacking pair of ger than maximally wide. First flagello- lateral carina entirely separated from me- mere 1.3 times longer than second dian bump. First tergite [0.8]-0.9 times flagellomere and 1.3 times longer than longer than wide. Border between first third flagellomere. Third flagellomere 1.5 and second tergite straight with edges times longer than wide. Flagellomere curving anteriorly. Second tergite smooth, length equal to or greater than width, with slightly elevated pinched-up area an- Horizontal length of eye 0.9 times longer teriorly. Suturiform articulation with than length of head behind eye. Trans- weak, smooth, V-shaped, weakly arched verse diameter of posterior ocellus 1.6 groove; with carina present along anterior times post-ocellar length. Distance be- margin. Second tergite [0.4]-0.5 times Ion- tween posterior ocellus and eye 5.3 times ger than wide, [0.8]-0.9 times longer than post-ocellar length. Longitudinal bump third tergite. Third tergite [0.5]-0.6 times between antennal sockets present. Facial longer than wide. Hypopygium with con- ridges bowed outward. Deep anterior vex dorsal margin. Ovipositor 1.1— [1.2] groove between antennal sockets absent, times body length. Area between ridges not filled in, level Color. — Black except as follows: maxil- with rest of face. Median carina on face lary and labial palpomeres white, metan- present, developed into raised tear-drop otum sometimes yellowish orange, pro- shaped area. Area between median carina podeum yellowish orange, fore tarsus yel- and ridges with ladder-like series of hor- lowish orange but telotarsus black, terga izontal carinae. Ridges running at 45° an- 1-4 yellowish orange. Fore wing banded gle from middle ridge to antennal sockets clear, black, clear, black; hind wing yellow strong. Groove around eyes present and in basal half, black apically. smooth. Eye height 1.3 times greater than Distribution and material examined. — eye width, 1.3 times greater than width of Known only from type locality in the Al- face. Width of head 2.4 times greater than 168 Journal of Hymenoptera Research width of face. Inter-tentorial distance 2.5 times clypeus height. Tentorio-ocular dis- tance 1.1 times longer than clypeus high. Clypeus partially filled in dorsally. Ven- tral margin of clypeus concave. Area around clypeus not differentiated from rest of face. Face 1.0 times wider than high. Malar suture paralleled by 8 ridges. Malar space 0.2 times eye height. Mesosoma. — Mesosoma 1.6 times longer than high. Pronotum with weak groove anteriorly. Notaulus not impressed. Bor- der between mesoscutum and scutellum with carinate groove, lacking median ca- rina and enlarged median pit. Propodeal spiracle crescent-shaped, 1.9 times higher than wide. Fore tibia 1.1 times longer than fore fe- mur. Fore tarsus not laterally compressed, 1.7 times longer than fore femur. Fore bas- itarsus 5.2 times longer than wide, 1.7 times longer than second tarsomere. Hind femur 4.1 times longer than wide, 1.5 times longer than basitarsus. Hind tibia 2.6 times longer than basitarsus. Outer and inner hind tibial spurs 0.3 and 0.5 times longer than basitarsus, respectively. Hind basitarsus 5.5 times longer than wide. Fore wing venation: 1M and lcu-a in- tersect. (RS + M)b broken apically. 1M 2.3 times longer than IRS. 2M 3.5 times longer then r-m. lm-cu 1.3 times longer than r, 1.7 times as thick as (RS + M)a. 2RS 1.2 times longer than r-m. 3RSa 2.9 times lon- ger than r-m, 4.8 times longer than r. 3RSb 5.4 times longer than r. C+SC + R and IRS forming an angle of 71°. Fore wing length 9.0 mm. Hind wing vein Rla 1.8 times longer than lr-m. Metasoma. — Base of petiole deeply ex- cavated. First tergite with strongly raised tongue-shaped bump. First tergite lacking lateral carina adjacent to median bump, with pair of lateral carina entirely sepa- rated from median bump. First tergite 1.1 times longer than wide. Border between first and second tergite straight with edges curving anteriorly. Second tergite smooth, with slightly elevated pinched-up area an- teriorly. Suturiform articulation barely distinguished from remainder of synter- gite, lacking groove, M-shaped, weakly arched; lacking carina along anterior mar- gin. Second tergite 0.7 times longer than wide 1.4 times longer than third tergite. Third tergite 0.5 times longer than wide. Hypopygium with convex dorsal border. Ovipositor 1.1 times body length. Color. — Yellowish orange and black. Head black with maxillary and labial pal- pomeres yellowish orange. Mesosoma yellowish orange except pro- pleuron black. Fore and mid legs yellow- ish orange except both coxae black and tarsi yellowish orange with telotarsus black. Hind leg black except some yellow- ish orange on basal part of tibia. Fore wing banded: black, yellow, black, yellow, black, costa black. Hind wing banded: yel- low, black, clear. Terga 1-4 yellowish orange. Terga 5-8 and ovipositor sheath black. Distribution and material examined. — Known only from type specimen. 1 9 ex- amined. Holotype 9. SURINAME: Suriname Exp. 1948- 1949, Nassau Mts., Malowijne, 15.ii.1949, D.C. Ge- ijskes (RMNH— JL000202). Remarks. — This species is very strange and may not be a member of Cyelaulacidea. Although it has the tear-drop shaped me- dian area of the face, a putative synapo- morphy for Cyelaulacidea, it lacks the well- developed suturiform articulation and rectangular bump on the first tergite that all other species of Cyelaulacidea possess. Etymology.— For John Pickering of The University of Georgia. Cyelaulacidea pottsae Leathers n. sp. Fig. 12B Diagnosis. — Fore wing black with one clear stripe. Hind wing black with clear apical tip. Terga 1-7 black dorsally, red- dish orange laterally (Fig. 12B). Volume 14, Number 2, 2005 169 Length. — 8.0 mm. Head. — Antenna with 46 flagellomeres. Scape longer ventrally than dorsally. Scape with shelf-like process apically, lacking pre-apical shelf, 1.5 times longer than maximally wide. First flagellomere 1.4 times longer than second flagellomere, 1.5 times longer than third flagellomere. Third flagellomere 1.1 times longer than wide. Apical flagellomere 1.7 times longer than wide. Flagellomere length equal to or greater than width. Horizontal length of eye 1.5 times longer than length of head behind eye. Transverse diameter of pos- terior ocellus 0.8 times post-ocellar length. Shortest distance between posterior ocel- lus and eye 2.9 times post-ocellar length. Longitudinal bump between antennal sockets present. Facial ridges diverging outward straight from clypeus to antennal sockets. Deep anterior groove between an- tennal sockets absent. Area between ridg- es not filled in, level with rest of face. Me- dian carina on face present, developed into raised tear-drop shaped area. Area between median carina and ridges with ladder-like series of horizontal carinae. Ridges running at 45° angle from middle ridge to antennal sockets strong. Groove around eyes present and crenulate. Eye height 1.3 times greater than eye width, 1.2 times greater than width of face. Width of head 2.3 times greater than width of face. Inter-tentorial distance 2.5 times clypeus height. Tentorio-ocular distance 1.3 times longer than clypeus high. Clyp- eus completely filled in but uneven. Ven- tral margin of clypeus concave. Area around clypeus with series of large cren- ulae and sharp ridge separating from rest of face. Face 1.1 times wider than high. Malar suture paralleled by 4 ridges. Malar space 0.2 times eye height. Mesosoma. — Mesosoma 1.5 times longer than high. Pronotum with deep, smooth groove anteriorly. Notaulus not im- pressed. Border between mesoscutum and scutellum with carinate groove, lacking median carina and median area not en- larged into a median pit. Propodeal spi- racle oval, 2.0 times higher than wide. Fore tibia 1.2 times longer than fore fe- mur. Fore tarsus not laterally compressed, 1.6 times longer than fore femur. Fore bas- itarsus 4.2 times longer than wide, 1.8 times longer than second tarsomere. Hind femur 3.8 times longer than wide, 1.8 times longer than basitarsus. Hind tibia 2.6 times longer than basitarsus. Outer and inner hind tibial spurs 0.4 and 0.6 times longer than basitarsus, respectively. Hind basitarsus 5.5 times longer than wide. Fore wing venation: 1M and leu-a in- tersect. (RS+M)b broken apically. 1M 2.1 times longer than IRS. 2M 3.6 times longer then r-m. lm-cu 1.1 times longer than r, 1.8 times as thick as (RS+M)a. 2RS 1.1 times length of r-m. 3RSa 3.2 times longer than r-m, 5.1 times longer than r. 3RSb 4.7 times longer than r. C + SC + R and IRS forming an angle of 78°. Fore wing length 7.6 mm. Hind wing vein Rla 1.5 times longer than lr-m. Metasoma. — Base of petiole deeply ex- cavated. First tergite with strongly raised rectangular bump. First tergite with lateral carina closely paralleling median bump but not forming notches, lacking pair of lateral carina entirely separated from me- dian bump. First tergite 0.9 times longer than wide. Border between first and sec- ond tergite straight with edges curving anteriorly. Second tergite with strongly pinched-up area not reaching third tergite. Suturiform articulation with weak, smooth, M-shaped, weakly arched groove; with carina present along anterior margin. Second tergite 0.5 times longer than wide, 0.7 times longer than third tergite. Third tergite 0.6 times longer than wide. Hypo- pygium with flat dorsal border. Oviposi- tor 1.0 times body length. Color. — Mostly black except maxillary and labial palpomeres yellowish orange apically, margins of eyes yellowish or- ange, and terga 1-7 reddish orange later- 170 Journal of Hymenoptera Research ally- Fore wing black with one clear stripe. Hind wing black with clear apical tip. Distribution and material examined. — Known only from type specimen. 19 ex- amined. Holotype ?. MEXICO: Orizaba, 1867, O. Sichel, (MNHN— JL000243). Etymology. — For Martha Potts of The University of Kentucky. Cyclaulacidea riceorum Leathers n. sp. Figs. 6B, 11B Diagnosis. — Fore wing veins 1M and lcu-a intersect. Hind tarsus black but bas- al part of basitarsus orange. Ovipositor 1.2 times body length. Horizontal length of eye (in dorsal view) 1.0-1.4 times longer than length of head behind eye. Fore tibia entirely yellowish orange (Fig. 11B). Length.— 7.8-[10.1] mm. Head.— Antenna with 48-[52] flagello- meres. Scape longer ventrally than dorsal- ly, with shelf-like process apically, lacking pre-apical shelf, 1.7-[2.1] times longer than maximally wide. First flagellomere 1.3-1.5 [1.4] times longer than second fla- gellomere and 1.4-[1.6] times longer than third flagellomere. Third flagellomere [1.2]-1.3 times longer than wide. Apical flagellomere [1.8J-2.9 times longer than wide. Flagellomere length equal to or greater than width. Horizontal length of eye [1.0]-1.4 times longer than length of head behind eye. Transverse diameter of posterior ocellus 0.9-[1.2] times post-ocel- lar length. Shortest distance between pos- terior ocellus and eye 2.8-[4.0] times post- ocellar length. Longitudinal bump be- tween antennal sockets present. Facial ridges parallel. Deep anterior groove be- tween antennal sockets very weak. Area between ridges not filled in, level with rest of face, or filled in creating a raised me- dian area. Median carina on face present, developed into raised tear-drop shaped area. Area between median carina and ridges with ladder-like series of horizontal carinae. Ridges running at 45° angle from middle ridge to antennal sockets strong. Groove around eyes present and crenu- late. Eye height [1.3]-1.4 times greater than eye width, 1.2-1.7 [1.3] times greater than width of face. Width of head 2.1-2.7 [2.2] times greater than width of face. In- ter-tentorial distance 1.9-3.1 [2.0] times clypeus height. Tentorio-ocular distance 0.7-1.1 [0.9] times longer than clypeus high. Clypeus partially filled in dorsally. Ventral margin of clypeus concave. Area around clypeus with series of large cren- ulae but lacking ridge, or [with series of large crenulae and sharp ridge separating from rest of face]. Face 0.7-1.1 [0.8] times wider than high. Malar suture paralleled by 1-4 [3] ridges. Malar space 0.1-[0.2] times eye height. Mesosoma. — Mesosoma 1.6-[1.7] times longer than high. Pronotum with deep, smooth groove anteriorly. Notaulus not impressed. Border between mesoscutum and scutellum with carinate groove. Bor- der between mesoscutum and scutellum lacking median carina and enlarged me- dian pit. Propodeal spiracle [oval] to cres- cent-shaped, 2.0-2.5 [2.2] times higher than wide. Fore tibia 1.1— [1.2] times longer than fore femur. Fore tarsus not laterally com- pressed, [1.6]-1.7 times longer than fore femur. Fore basitarsus 4.3-5.4 [5.2] times longer than wide, 1.5-1.8 [1.6] times lon- ger than second tarsomere. Hind femur 3.8-[4.0] times longer than wide, [1.6]-1.8 times longer than basitarsus. Hind tibia [2.3]-2.7 times longer than basitarsus. Outer and inner hind tibial spurs [0.4]-0.5 and 0.6 times longer than basitarsus, re- spectively. Hind basitarsus 5.0-6.2 [6.0] times longer than wide. Fore wing venation: 1M and lcu-a in- tersect. (RS+M)b broken apically. 1M 1.9- 2.1 [2.0] times longer than IRS. 2M 3.3- [3.8] times longer then r-m. lm-cu 1.0-1.5 [1.2] times longer than r, 1.1-2.2 [1.5] times as thick as (RS+M)a. 2RS 1.2-[1.3] times longer than r-m. 3RSa 2.7-[3.2] times lon- ger than r-m, 4.3-6.2 [5.0] times longer Volume 14, Number 2, 2005 171 than r. 3RSb [5.2]-7.2 times longer than r. mostly black with some yellowish orange C + SC + R and IRS forming an angle of 72- by trochanter. Hind tibia black, or mostly 78° [74°]. Fore wing length 7.3-[8.6] mm. black but yellowish orange basally. Hind Hind wing vein Rla [1.4]-1.6 times Ion- tarsus black but basal part of basitarsus ger than lr-m. orange. Fore wing banded: yellow, black, Metasoma. — Base of petiole deeply ex- yellow, black, costa yellow. Hind wing cavated. First tergite with strongly raised yellow in basal half, black apically, or rectangular bump. First tergite with lateral banded: yellow, black, yellow, carina closely paralleling median bump Terga 1-3 yellowish orange. Tergum 4 but not forming notches, lacking pair of yellowish orange anteriorly, black poste- lateral carina entirely separated from me- riorly. Tergum 5 black, or yellowish or- dian bump. First tergite 0.8-1.1 [1.0] times ange anteriorly, black posteriorly. Terga longer than wide. Border between first 6-8 and ovipositor sheath black, and second tergite straight with edges Distribution and material examined. — curving anteriorly. Second tergite smooth, Known from Peru and Brazil. 29 9, 46*6* with slightly elevated pinched-up area an- examined. teriorly. Suturiform articulation with Holotype ?. PERU: Madre de Dios: Rio Tambo- weak, smooth, V-shaped, weakly arched pata Reserve, 30km (air) SW Puerto Maldonado, groove; with carina along anterior margin. 290m, 25-30.iv.l984, W.J. Pulawski (CAS— JL000094). Second tergite [0.5]-0.6 times longer than Paratypes. PERU: 1 ?, Madre de Dios: Rio Tambopata •j n n n n rn on i-- i j.u „ u,- a Reserve, 30km (air) SW Puerto Maldonado, 290m, 25- wide, 0.7-0.9 [0.8] times longer than third ' f.r/ ' .. fr.AC TTnnmA~, », , r_ J? 30. iv. 1984, W.J. Pulawski (CAS — JL000162); Monzon tergite. Third tergite 0.6-[0.7] times longer Valley/ Tingo Mari3/ EJ Schlinger & E.s. Ross: 16 than wide. Hypopygium with convex dor- 2.xi.l954 (CAS— JL000240), 13 19. x. 1954 (CAS- sal border. Ovipositor 1.2 times body JL000241), 13 3.xi.l954 (CAS— JL000242). BRAZIL: length *^d ' Utiariti (325m), Rio Papagaio, viii.1961, K. Lenko Co/or.-Black and yellow. Head black (MZSP-JL000244). except maxillary and labial palpomeres Etymology. — For Harold and Leona Rice and sometimes areas around margins of for their generous support of this research eyes and posterior to malar suture yellow- and systematic entomology at Oregon ish orange. State University. Mesosoma black except sometimes dor- sal parts of propodeum yellowish orange. Cyclaulacidea rominus Leathers n. sp. Forecoxa, trochanter, trochantellus, and Hgs- ^K' lilJ femur black or yellowish orange. Fore tib- Diagnosis. — Fore wing banded yellow, ia yellowish orange. Fore tarsus yellowish black, yellow, black; costa black (Fig. 11D). orange but telotarsus black. Mid coxa Mid femur entirely black. Facial ridges black, or yellowish orange, or mostly yel- bowed outward (Fig. 2B). Propodeal spi- lowish orange with some black basally. racle crescent-shaped, [2.7J-3.2 times high- Mid trochanter black, or yellowish orange, er than wide. Mid trochantellus yellowish orange. Mid Length. — [8.8]-9.6 mm. femur black, or yellowish orange, or most- Head. — -Antenna with [48]-53 flagello- ly black, with some yellowish orange on meres. Scape longer ventrally than dorsal- both ends. Mid tibia yellowish orange, or ly. Scape without any shelf-like process yellowish orange but black in apical quar- apically, or with shelf-like process apical- ter. Mid tarsus mostly yellowish orange ly, lacking pre-apical shelf. Scape 1.6-[1.7] but telotarsus black. Hind coxa and tro- times longer than maximally wide. First chanter black. Hind trochantellus yellow- flagellomere [1.2J-1.4 times longer than ish orange. Hind femur black, or black in second flagellomere, [1.3]-1.4 times longer apical half, yellowish orange basally, or than third flagellomere. Third flagello- 172 Journal of Hymenoptera Research mere 1.0— [1.1] times longer than wide. Apical flagellomere 1.8-[1.9] times longer than wide. Flagellomere length [equal to or greater than width], or distinctly less than width. Horizontal length of eye 1.4- [1.7] times longer than length of head be- hind eye. Transverse diameter of posterior ocellus 1.0— [1.1] times post-ocellar length. Distance between posterior ocellus and eye 3.5 times post-ocellar length. Longi- tudinal bump between antennal sockets present. Facial ridges bowed outward. Deep anterior groove between antennal sockets very weak. Area between ridges not filled in, level with rest of face. Median carina on face present, developed into raised tear-drop shaped area. Area be- tween median carina and ridges with lad- der-like series of horizontal carinae. Ridg- es running at 45° angle from middle ridge to antennal sockets strong. Groove around eyes present and crenulate. Eye height 1.3 times greater than eye width, 1.2 times greater than width of face. Width of head 2.2 times greater than width of face. Inter- tentorial distance 2.0-[2.3] times clypeus height. Tentorio-ocular distance 1.0— [1.1] times longer than clypeus high. Clypeus partially filled in dorsally. Ventral margin of clypeus concave. Area around clypeus with series of large crenulae and sharp ridge separating from rest of face. Face 1.1— [1.2] times wider than high. Malar su- ture paralleled by 2 ridges. Malar space 0.2 times eye height. Mesosoma. — Mesosoma [1.4]-1.5 times longer than high. Pronotum with deep, smooth groove anteriorly. Notaulus not impressed. Border between mesoscutum and scutellum lacking groove, or [with carinate groove]. Border between mesos- cutum and scutellum lacking median ca- rina and not enlarged into a median pit. Propodeal spiracle crescent-shaped, [2.7]- 3.2 times higher than wide. Fore tibia [1.1]-1.2 times longer than fore femur. Fore tarsus not laterally com- pressed, [1.6]-1.7 times longer than fore femur. Fore basi tarsus [3.3]-4.3 times lon- ger than wide, [1.6]-1.7 times longer than second tarsomere. Hind femur 3.7-[4.1] times longer than wide, 1.9-[2.1] times longer than basitarsus. Hind tibia [2.5]-2.6 times longer than basitarsus. Outer and inner hind tibial spurs 0.5 and [0.6]-0.7 times longer than basitarsus, respectively. Hind basitarsus [4.2J-4.5 times longer than wide. Fore wing venation: 1M and lcu-a in- tersect. (RS+M)b broken apically. 1M 1.5- [1.7] times longer than IRS. 2M 3.3-L3.8] times longer then r-m. lm-cu 1.0 times longer than r, 1.5-[2.0] times as thick as (RS + M)a. 2RS 1.0-[1.2] times longer than r-m. 3RSa 2.8-[3.1] times longer than r-m, [4.0]-4.4 times longer than r. 3RSb 4.6- [5.0] times longer than r. C + SC + R and IRS forming an angle of 73°. Fore wing length [9.0J-10.3 mm. Hind wing vein Rla 1.4 times longer than lr-m. Metasoma. — Base of petiole deeply ex- cavated. First tergite with strongly raised rectangular bump. First tergite with lateral carina closely paralleling median bump but not forming notches, lacking pair of lateral carina entirely separated from me- dian bump. First tergite 0.9-[1.0] times longer than wide. Border between first and second tergite [straight with edges curving anteriorly], or completely round- ed. Second tergite with strongly pinched- up area not reaching third tergite. Suturi- form articulation with weak, smooth, V- shaped, weakly arched groove; with cari- na along anterior margin. Second tergite 0.4-[0.5] times longer than wide, 0.7-[0.9] times longer than third tergite. Third ter- gite [0.5]-0.6 times longer than wide. Hy- popygium with flat to [convex] dorsal bor- der. Ovipositor 0.9 times body length. Color. — Head black except maxillary and labial palpomeres black basally, yel- lowish orange apically, or entirely yellow- ish orange. Mesopleuron mostly black. Metanotum yellowish orange. Propodeum yellowish orange, or black laterally, yellowish or- Volume 14, Number 2, 2005 173 ange dorsally. Metapleuron black, or mostly black with some yellowish orange by hind wing. Legs entirely black except as follows: fore tibia black but yellowish orange in basal twelfth, or black but yel- lowish orange in basal and apical sixths. Fore tarsus yellowish orange but telotar- sus black. Mid trochantellus yellowish or- ange. Mid tibia mostly black, but basal fifth yellowish orange. Mid tarsus mostly yellowish orange but telotarsus black. Hind trochantellus yellowish orange. Hind tibia mostly black but yellowish or- ange basally. Hind tarsus mostly black but basal part of basitarsus and 4th tarsomere orange, or black but basal part of basitar- sus orange. Fore wing banded: yellow, black, yellow, black, costa black. Hind wing banded: yellow, black, yellow, black apical black band complete, or banded: yellow, black, yellow. Terga 1-4 yellowish orange. Tergum 5 black or yellowish orange. Terga 6-8 and ovipositor sheath black. Distribution and material examined. — Known only from Suriname. 29 9 exam- ined. Holotype 9. SURINAME: Boven Corantyn, Coer- oeni eil., 7.x. 1959, Creutzberg, (RMNH— JL000228). Paratype. SURINAME: 19, Tapanahoni Saniki, 9.V.1954, D.C. Geijskes (RMNH— JL000229). Etymology. — For Michael Robertson. Cyclaulacidea sharkeyi Leathers n. sp. Figs. 3B,'l2A Diagnosis. — Ovipositor sheath black with some yellowish orange in apical third (but black at apical tip) (Fig. 12A). Suturiform articulation with shallow groove (as in Fig. 7B). Strongly raised bump on petiole rectangular (as in Fig. 8B). Length.— 8.8-[14.1] mm. Head.— Antenna with 54-58 [57J flagel- lomeres. Scape longer ventrally than dor- sally. Scape with shelf-like process apical- ly, lacking pre-apical shelf, 1.9-2.3 [2.2] times longer than maximally wide. First flagellomere 1.2-1.5 [1.4] times longer than second flagellomere, 1.4-[1.6] times longer than third flagellomere. Third fla- gellomere [1.1]— 1.3 times longer than wide. Apical flagellomere 1.8-2.3 [2.1] times longer than wide. Flagellomere length [equal to or greater than width], or distinctly less than width. Horizontal length of eye [0.7]-0.8 times longer than length of head behind eye. Transverse di- ameter of posterior ocellus 0.9-1.4 [1.1] times post-ocellar length. Distance be- tween posterior ocellus and eye 3.8-5.6 [4.8] times post-ocellar length. Longitudi- nal bump between antennal sockets pre- sent. Facial ridges [parallel] or diverging outward straight from clypeus to antennal sockets. Deep anterior groove between an- tennal sockets very weak. Area between ridges filled-in creating a raised median area. Median carina on face present; de- veloped into raised chevron or butterfly- shaped area. Area between median carina and ridges smooth. Ridges running at 45° angle from middle ridge to antennal sock- ets absent, or weak, or [strong]. Groove around eyes present and smooth. Eye height 1.3-1.5 [1.4] times greater than eye width; 1.0-1.2 [1.1] times greater than width of face. Width of head 1.9-2.3 [2.1] times greater than width of face. Inter-ten- torial distance [1.7]-2.3 times clypeus height. Tentorio-ocular distance 1.0-1.3 [1.2] times longer than clypeus high. Clyp- eus completely filled in but uneven. Ven- tral margin of clypeus [flat], or concave. Area around clypeus with series of large crenulae and sharp ridge separating from rest of face. Face [1.1]— 1.2 times wider than high. Malar suture paralleled by [2]- 5 ridges. Malar space [0.2]-0.3 times eye height. Mesosoma. — Mesosoma 1.4-1.7 [1.5] times longer than high. Pronotum with deep, smooth groove anteriorly. Notaulus not impressed, or [weakly impressed]. Border between mesoscutum and scutel- lum with carinate groove. Border between mesoscutum and scutellum lacking medi- an carina; median area not enlarged, or 174 Journal of Hymenoptera Research enlarged into a median pit. Propodeal spi- racle oval, 1.8-[2.4] times higher than wide. Fore tibia 1.0-[1.2] times longer than fore femur. Fore tarsus not laterally com- pressed, 1.5-1.7 [1.6] times longer than fore femur. Fore basitarsus 3.9-5.7 [4.3] times longer than wide, [1.7]-1.9 times longer than second tarsomere. Hind femur 3.8-[4.2] times longer than wide, 1.6-[1.8] times longer than basitarsus. Hind tibia [2.5] -2.6 times longer than basitarsus. Outer and inner hind tibial spurs 0.4 and 0.5-[0.6] times longer than hind basitarsus. Hind basitarsus 5.2-6.2 [6.1] times longer than wide. Fore wing venation: [1M and lcu-a in- tersect], or lcu-a intersects Cu distad 1M. (RS+M)b completely tubular, or broken apically. 1M 1.8-2.1 [1.9] times longer than IRS. 2M [3.1]-3.6 times longer then r-m. lm-cu [1.2]-1.6 times longer than r, 1.1- 1.7 [1.6] times as thick as (RS+M)a. 2RS [1.0]-1.2 times longer than r-m. 3RSa [2.9]- 3.2 times longer than r-m, [5.0]-6.3 times longer than r. 3RSb [4.6]-6.0 times longer than r. C+SC + R and IRS forming an an- gle of 67-73° [72°]. Fore wing length 8.7- [12.8] mm. Hind wing vein Rla [1.3]-1.5 times lon- ger than lr-m. Metasoma. — Base of petiole not deeply excavated. First tergite with strongly raised rectangular bump. First tergite with lateral carina closely paralleling median bump but not forming notches, lacking pair of lateral carina entirely separated from median bump. First tergite 1.1— [1.3] times longer than wide. Border between first and second tergite straight with edges curving anteriorly. Second median tergite smooth, with slightly elevated pinched-up area anteriorly. Suturiform articulation with weak, smooth, M-shaped, weakly arched groove; with carina along anterior margin. Second tergite [0.5]-0.6 times lon- ger than wide, 0.7-0.9 [0.8] times longer than third tergite. Third tergite [0.5]-0.7 es longer than wide. Hypopygium with convex dorsal border. Ovipositor 0.8-1.1 [1.0] times body length. Color. — Yellowish orange and black. Head black except maxillary and labial palpomeres yellowish orange. Mesosoma yellowish orange except pro- pleuron black. Fore and mid legs yellow- ish orange except mid tibia sometimes black in apical quarter and telotarsus of mid tarsus black. Hind coxa black. Hind trochanter black, or yellowish orange. Hind trochantellus yellowish orange. Hind femur yellowish orange in basal half, black in apical half, or mostly black but yellowish orange in basal twelfth. Hind tibia mostly black but yellowish or- ange basally. Hind tarsus black. Fore wing banded: yellow, black, yellow, black, costa yellow. Hind wing yellow in basal half, black apically. Terga 1-4 yellowish orange. Tergum 5 yellowish orange, or yellowish orange an- teriorly, black posteriorly, or black dorsal- ly, yellowish orange laterally. Tergum 6 black, or black dorsally, yellowish orange laterally, or mostly yellowish orange with some black spots anteriorly. Tergum 7 black, or black dorsally, yellowish orange laterally. Tergum 8 black. Ovipositor sheath black with some yellowish orange in apical third (but black at tip). Distribution and material examined. — Known only from type locality of Dirk- shoop, Suriname. 49 9, 16 examined. Holotype $. SURINAME: Dirkshoop, 21. v. 1963, J.v.d. Vecht, (RMNH— JL000203). Paratypes. SURI- NAME: Dirkshoop, 21.V.1963, J.v.d. Vecht: 39? (RMNH— JL000236, 000238, 000239), 120% on claustral colonies of C. nigri- mal tergite of the female, which has strong ceps (n = 511), and from <5% to 17.5% on submedial posterior emarginations that C. mimosae (n = 475) (Stanton et al. in define a medial and pair of sublateral press). Parasitism by T. laikipiensis sp. nov. points. The genus may be identified using 184 Journal of Hymenoptera Research -vJ'r' Fig. 1. Trigastrotheca laikipiensis Quicke sp. nov., life stages: a, mid instar larva feeding on Cremastogaster nigriceps eggs with foundress queen in lower right; b, T. laikipiensis sp. nov. cocoon in situ protected by C. nigriceps foundress queen. Volume 14, Number 2, 2005 185 Fig. 2. Late instar larva of Trigastrotheca laikipiensis Quicke sp. nov., feeding on well-developed Cremas- togaster nigriceps larva. the key to the Old World genera of Bra- coninae of Quicke (1987). The genus Kenema was erected based on a single male specimen from Sierra Leone (van Achterberg 1983), and subsequently a 2nd species from Senegal was described by van Achterberg and Sigwalt (1987). Ac- cording to their diagnostic key, published in the same year that Quicke (1987) syn- onymized Kenema with Trigastrotheca, Ke- nema differs from the latter in that the 5th metasomal tergite completely lacks an ap- icomedial protuberance and is longer compared to the 4th; it also has less well developed anterolateral grooves on ter- gites 3-5, and fore wing vein r originates slightly more distally on the pterostigma. Based on our examination of much mate- rial in various collections, and addition- ally members of both sexes of the new species of Trigastrotheca described below, it is clear that the posterior margin of the 5th metasomal tergite is not always triden- tate in Trigastrotheca. In males it is simple, though relatively a little longer than in most other genera, and in the case of the new species described here, can have a small sub-posterior medial hump that is presumably a reduced expression of the female's strong medial point. Further, the finely serrate apparent posterior margin of the 4th and 5lh metasomal tergites pro- duced by an overhang of the coriaceous sculptured dorsal surface over the true margin in the new species is closely sim- ilar to that in Kenema serrata van Achter- berg & Sigwalt, suggesting that this might be a synapomorphy for the two species. Because of these observations, the synon- ymy of Kenema with Trigastrotheca origi- nally proposed by Quicke (1987) is hereby supported. Trigastrotheca laikipiensis Quicke, new species (Figs 2-4) Holotype female. — Laikipia District, Kenya. Collect- ed 11 July 2001 at Mpala Research Centre by M. Stan- ton. Found sealed within a swollen thorn of Acacia drepanolobium with a claustral colony of Crematogaster nigriceps ants. Bohart Museum of Entomogy; Univer- sity of California; Davis, CA USA. Paratypes. — 2 9 9 and 1 8: Laikipia District, Kenya. Collected 20 July 2001 at Mpala Research Centre by M. Stanton. Found sealed within swollen thorns of Acacia drepanolobium occupied by claustral colonies of Crematogaster mimosae ants. BMNH. Length of body, 4.3 mm, forewing 3.7mm, antenna 2.7mm and part of ovi- positor exserted beyond apex of metaso- ma 1mm. Head strongly transverse, densely se- tose; more or less uniformly coriaceous- rugulose. Antenna robust, with 26-27 fla- gellomeres. Terminal flagellomere robust and distinctly acuminate. Median flagel- lomeres marginally longer than wide. Height of clypeus: inter-tentorial distance: tentorio-ocular distance = 1.0:3.6:2.2. Face with distinct mid-longitudinal carina on somewhat protruding upper third. Width of head: height of eye: shortest distance between eyes = 2.8:1.0:1.6. Eyes with short sparse setae. Frons with strong mid-lon- gitudinal ridge ending abruptly shortly before anterior ocellus. Posterior ocellar line : transverse diameter of posterior ocel- lus: shortest distance between posterior ocellus and eye = 4.4:1.0:3.7. Mesosoma almost entirely coriaceous. Notauli well developed; mesoscutum with a weak but distinct mid-longitudinal ca- 186 Journal of Hymenoptera Research Fig. 3. Automontage® habitus pictures of Trigastrotheca laikipiensis Quicke sp. nov., female holotype: a, lateral aspect, b, dorsal aspect. rina. Pronotum laterally with a deep cren- ulated curved groove and with strong sub-longitudinal striation below this. Me- sopleuron with a weakly depressed broad mesopleural suture, and with an appar- ently weakly sclerotized almost complete rnaulus below this. Median area of me- tanotum with complete mid-longitudinal carina. Propodeum with a complete mid- longitudinal (sub-lamelliform) carina run- ning within a distinct crenulated groove; with distinct straight sub-lateral carinae ending at level of spiracle just above a dis- tinct strongly setose protuberance; with Volume 14, Number 2, 2005 187 curved ('C'-shaped) crenulated groove be- low spiracle; posteriorly with a few dis- tinct anteriorly-diverging short carinae. Claws with small but distinctly pointed basal lobes. Lengths of fore femur : tibia : tarsus = 1.0:1.0:1.14. Lengths of hind fe- mur: tibia : tarsus = 1.0:1.4:1.3. Hind fe- mur very robust, 3.5 times longer than maximally deep. Forewing veins 1-M and 1-SR + M meet- ing at an acute angle near wing margin, vein 1-SR short, forming an angle of 85° with C + SC + R. Lengths of veins r:3-SR: SRI = 1.0:1.36:5.0. Vein r arising 0.28-0.3 from the base of pterostigma. Pterostigma 3.2 times longer than broad. Lengths of veins 2-SR : 3-SR : r-m = 1.45:1.75:1.0. Vein 2-SR+M slightly longer than lm-cu. Vein 3-CU1 forming a continuous curve with CUlb. Metasoma strongly coriaceous. Tergites 3-5 with well developed antero-lateral tri- angular areas. Posterior margins of 3rd and 4th metasomal tergites appearing strongly dentate, but this is due to the strong co- riaceous-rugose sculpture combined with a steep posterior margin such that the in- dividual confluences of rugae, each of which bears a posteriorly-directed seta, overhangs the true posterior tergal mar- gins. Posterior margin of 5th tergite of fe- male with deep sub-median semicircular emarginations creating sharp medial point between them; lateral to emarginations border with fine serration. Ovipositor lacking pre-apical dorsal nodus and with no lower valve teeth visible at X40; in lat- eral aspect strongly depressed at apex. Coloration mostly ochreous yellow with variable dark markings (mark on top of head formed into two points anteriorly, large marks on the medial and lateral lobes of the mesoscutum, mesosternum and metasomal sternites), and with whit- ish, less sclerotized, longitudinal line along the position of the sternaulus (i.e. below the depressed part of the meso- pleural suture). Antennae black; mandi- bles whitish with blackish tips; wing membrane weakly greyish, wing venation dark brown to blackish. Ovipositor pale brownish. Variation. — Some paratypes are slightly less extensively marked with black or dark brown, one female paratype having dark spots antero-laterally on the propo- deum. The male has the propodeum, scu- tellum and most of the mesoscutum ex- cept a small medio-posterior part, a mark on the middle of the 1st metasomal tergite, and small spots mediolaterally on the 3rd and 4th tergites blackish. Notes. — Nine species of Trigastrotlieca have been described previously: T. inermis (Guerin-Meneville) comb. nov. (on au- thority of C. van Achterberg) [Spinaria inermis Guerin-Meneville, 1848], T. nigri- cornis Cameron, T. quickei (van Achter- berg), T. romani nom. nov. [= Coelodontus costator: Roman; = Ichneumon costator Thunberg a junior homonym of Ichneumon costator Donovan, 1810; see Yu & Horst- man (1997)], T. rugosa (Szepligeti) comb, nov. [= Bracon rugosa Szepligeti; = K. ru- gosa: Quicke & Koch (1990)], T. serrata (van Achterberg & Sigwalt), T. tricolor Quicke & Ingram, T. tridentata (Enderlein) and T. trilobata Cameron. The new species described here can be distinguished from all other described species of Trigastrotheca by its smaller size and colour pattern. The apparently close- ly-related T. serrata differs further in hav- ing the frons weakly convex, distinctly striate and without a mid-longitudinal ca- rina, the mesoscutum with hardly any trace of notauli and without a mid-longi- tudinal ridge, the pronotum, propodeum and mesopleuron less heavily sculptured and the latter without mesopleural suture and sternaulus. DISCUSSION Parasites are known to have significant impacts on mature ant colonies in some systems (Morrison 1999, Feener 2000), but is not clear how common or ecologically important brood parasites of ant foun- 188 Journal of Hymenoptera Research Ki Fig. 4. Automontage® pictures of Trigastrotheca laikipiensis Quicke sp. nov., female holotype: a, head, anterior aspect; b, head, dorsal aspect; c, scutellum, propodeum and 1st two metasomal tergites; d, mesosoma, lateral aspect. dresses might be. We have never observed T. laikipiensis within swollen thorns on trees occupied by mature colonies, sug- gesting that this parasite specializes on the brood of young foundress queens (also see Yu and Quicke 1997). Many of the important elements of the host-parasite interaction described here re- main obscure. The biology of Trigastrothe- ca has previously been unknown. Mor- phologically (presence of complete mid- longitudinal propodeal carina) the genus appears to be related to several that are associated with galls, such as Simplicibra- con Quicke, Testudobracon Quicke and Triaspidogastra Granger. If this is the case then the group may historically be asso- ciated with distinct swellings on plant tis- sue, though in the other cases, oviposition appears to be through the plant tissue, and the ovipositor is typically furnished with a distinct dorsal nodus and apico- ventral serrations (Quicke 1987). We do not know how eggs of T. laikipiensis enter the sealed chambers of claustral colonies of acacia-ants on A. drepanolobium, but it is likely that female wasps locate swollen thorns with foundress queens inside, and then pierce the thorn wall or the queen's entry seal to deposit an egg within the claustral chamber. Within the Braconidae, parasitism of ants is rare, with only two previously known examples of its evolution. Mem- bers of the Neoneurus Haliday group of genera in the Euphorinae (previously treated as the subfamily Neoneurinae, but see Belshaw et al. 2001) are endoparasi- Volume 14, Number 2, 2005 189 Fig. 5. Automontage E" picture of Trigastrotheca laikipiensis Quicke sp. nov., female holotype: detail of posterior of metasoma, showing that dentation at posterior of 4th tergite is due to overhanging and seta-bearing sculp- turation. toids of adult worker ants (Shaw 1993). One species of the New World braconine genus Compsdbraconoid.es Quicke parasit- izes foundress queens of Aztecn ants living in domatia in the plant Cordia nodosa Lam. (Boraginaceae) (Yu and Quicke 1997), but other members of this group have more typical biologies for the subfamily (Fortier and Nishida 2004, D. Janzen, unpublished data). Given that the Trigastrotlieca wasps probably gain protection from their foun- dress hosts through chemical crypsis or mimicry, it would be interesting to know whether there is any differentiation be- tween individuals attacking the three ant species at our study sites. However, apart from some slight variation in coloration, all members of the individuals seen are morphologically very uniform, and there- fore we currently consider them to repre- sent a single species. ACKNOWLEDGEMENTS John Lemboi and Monica Turner provided expert field and lab assistance on this project, and Steve Heydon and Lynn Kimsev at the Bohart Museum graciously helped with preliminary wasp identifica- tion. Kees van Achterberg kindly allowed us to pub- lish on his identification of Spinaria inermis and ad- vised on the status of Coelodontus costator. Andy Po- laszek kindly provided Automontage' facilities. As always, the staff at the Mpala Research Centre pro- vided excellent logistical support. This work was sup- ported by NSF grants DEB-9726663 and DEB-0089706 to M. Stanton, T. Palmer, and T. P. Young. LITERATURE CITED Belshaw, R., C. Lopez- Vaamonde, N. Degerli, jnd I). L. J. Quicke. 2001. Paraplu letic taxa and taxo- nomic chaining: evaluating the classification ot braconine wasps (Hymenoptera: Braconidae) us- ing 28S D2-3 rDNA sequences and morpholog- ical characters. Biological Journal of the Linnear ciety 73: 411-424. Donovan, E. 1810. The Natural History of British In- sects. Vol. XIV. London. 90 pp. I eener, D. J. 2000. Is the assembly of ant communities mediated by parasitoids? Oikos 90: 79-88 Fortier, J. and K. Nishida. 2004. A new species and host association biolog) of Neotropical Compso braconoides Quicke (Hymenoptera: Braconidae). Journal of Hymenoptera Research 13: 228-233. Guerin-Meneville, F. E. 1848. Ordre des Hymenop- teres. In: Voyage en Abyssinie execute pendant 190 Journal of Hymenoptera Research les annees 1839-1843 (von Mr. Lefebvre). Ber- trand, Paris. Hocking, B. 1970. Insect associations with the swollen thorn acacias. Transactions of the Royal Entomolog- ical Society of London 122: 211-255. Morrison, L. W. 1999. Indirect effects of phorid fly parasitoids on the mechanisms of interspecific competition among ants. Oecologia (Berlin) 121: 113-122. Palmer, T. M. 2003. Spatial habitat heterogeneity in- fluences competition and coexistence in an Afri- can acacia ant guild. Ecology 84: 2843-2855. Palmer, T. M., M. L. Stanton, and T. P. Young. 2003. Competition and coexistence: Exploring mecha- nisms that restrict and maintain diversity within mutualist guilds. American Naturalist 162: S63- S79. Palmer, T. M., T. P. Young, M. L. Stanton, and E. Wenk. 2000. Short-term dynamics of an acacia ant community in Laikipia, Kenya. Oecologia (Berlin) 123: 425-435. Quicke, D. L. J. 1987. The Old World genera of bra- conine wasps (Hymenoptera: Braconidae). Jour- nal of Natural History 21: 43-157. Quicke, D. L. J. and S. N. Ingram. 1993. Braconine wasps of Australia. Memoirs of the Queensland Museum 33: 299-336. Quicke, D.L.J, and F. Koch. 1990. Die Braconinae-Ty- pen der beiden bedeutendsten Hymenopteren Sammlungen der DDR. Deutsches Entomologische Zeitschrift. 37: 213-227. Shaw, S. R. 1993. Observations on the ovipositional behavior of Neoneurus mantis, an ant-associated parasitoid from Wyoming (Hymenoptera: Bra- conidae). Journal of Insect Behavior 6: 649-658. Stanton, M. L., T. M. Palmer, and T. P. Young. 2002. Competition-colonization trade-offs in a guild of African Acacia-ants. Ecological Monographs 72: 347-363. . in press. Ecological barriers to early colony establishment in three coexisting acacia-ant spe- cies in Kenya. Insectes Sociaux. Taiti, S. W. 1992. The vegetation of Laikipia District Kenya. Laikipia- Mount Kenya Papers, B-2, Uni- versity of Bern; University of Nairobi. Thunberg, C.P. 1822. Ichneumonidea, Insecta Hyme- noptera illustrata. Memoires de VAcademie Imperi- ale des Sciences de Saint Petersbourg 8: 249-281. van Achterberg, C. 1983. Six new genera of Braconina from the Afrotropical Region (Hymenoptera, Braconidae). Tijdschrift voor Entomologie 126: 175- 202. van Achterberg, C. and B. Sigwalt. 1987. Three new genera of Braconinae from the Afrotropical re- gion (Hymenoptera: Braconidae). Zoologische Me- dedelingen 61: 443^58. Young, T. P., B. D. Okello, D. Kinyua, and T. M. Palmer. 1998. KLEE: a long-term multi-species herbivore exclusion experiment in Laikipia, Ken- ya. African Journal of Range Forage Science 14: 94- 102. Young, T. P., C. H. Stubblefield, and L. A. Isbell. 1997. Ants on swollen-thorn acacias: Species coexis- tence in a simple system. Oecologia (Berlin) 109: 98-107. Yu, D. S. and K. Horstmann. 1997. A catalogue of world Ichneumonidae (Hymenoptera). Memoirs of the American Entomological Institute, 58: 1-1558. Yu, D. W. and D. L. J. Quicke. 1997. Compsobraconoides (Braconidae: Braconinae), the first hymenopteran ectoparasitoid of adult Azteca ants (Hymenop- tera: Formicidae). Journal of Hymenoptera Research 6: 419-421. I. HYM. RES. Vol. 14(2), 2005, pp. 191-199 Pappognatha Mickel (Hymenoptera: Mutillidae: Sphaeropthalminae): New Species, Sex Associations, Hosts, and New Distribution Records DlOMEDES QUINTERO A.* AND ROBERTO A. CAMBRA T. Museo de Invertebrados G. B. Fairchild, Estafeta Universitaria 0824, Universidad de Panama, Panama, Rep. de Panama. *Author for correspondence: dquinter@ancon.up.ac.pa Abstract. — Two new species of Pappognatha are described: P. panamensis Quintero and Cambra, male and female, from Panama, and P. peruana Quintero and Cambra, male, from Peru. The previously unknown males of Pappognatha myrmiciformis (Cameron, 1897) and Pappognatha patruelis (Andre, 1898) are described, bodies and genitalia are illustrated. Pappognatha patruelis is reported as a parasitoid of Euglossa sp. (Apidae). New country records are presented for the following four species: P. speciosa Mickel, 1939, Colombia and Ecuador, previously known from Peru; P. egregia Mickel, 1939, Colombia and Ecuador, previously known from Brazil; P. obliqua Mickel, 1939, Peru, previously known from Brazil; P. myrmiciformis, Honduras, previously known from Panama and Costa Rica. Honduras represents the northernmost distribution record for this Neotropical genus. Pappognatha Mickel, 1939, is a Neotrop- 7) in both sexes of Pappognatha is probably ical genus of mutillid wasps, erected to an adaptation to tear open and exit the comprise 13 species based on single-sex hard resin nests of Euglossa by the newly descriptions: 11 known only from females eclosed adults. and 2 from males. Males of Pappognatha Because of the great resemblance be- are rare in collections, a total of five spec- tween P. myrmiciformis and the ant Cam- imens having been reported in the litera- ponotus sericeiventris Guerin-Meneville ture: two of P. obliqua Mickel, 1939, one of some authors (e.g. Wheeler 1983) assumed P. carmo (Cresson, 1902), and two of Pap- erroneously that the mutillid was a para- pognatha sp. (Quintero and Cambra 1996). sitoid or a brood parasite of that ant Females are more common, but relatively (Brothers et ah 2000). scarce in reference collections; only 50 fe- Mickel (1939) suggested two possible male specimens with locality records have sex associations, based on distribution and been cited in the literature (Mickel 1939, collection data: P. carmo (Cresson), proba- Cambra and Quintero 1992, Yanega 1994, bly the male of P. Uinta (Mickel), from San- Quintero and Cambra 1996). Pappognatha tarem, Brazil; and P. obliqua Mickel, prob- larvae parasitize immature Euglossa bees ably the male of P. pertyi (Dalla Torre), (Roubik 1989). Yanega (1994) reports two from Para, Brazil. findings of Pappognatha myrmiciformis as a Here we describe two new species oi parasitoid of Euglossa dodsoni Moure in Pappognatha; one of them being the first to Costa Rica and one record of Pappognatha be described from both sexes. We also de- speciosa Mickel as a parasitoid of Euglossa scribe the previously unknown males of P. intersecta Latreille that had nested inside patruelis (Andre, 1898) and P. myrmicifor- an aerial termite nest in Iquitos, Peru. No mis (Cameron, 1897), and provide an illus- additional hosts are known for Pappogua- trated key to the six species known from tha and apparently their parasitism is re- males. We present four new country re- stricted to bees of the genus Euglossa. The cords and the first host record for P. pa- development of strong mandibles (Figs. 5- truclis. We have examined new material of 192 Journal of Hymenoptera Research Pappognatha (46 females and 8 males), bringing the total of currently recorded Pappognatha to 94 females and 13 males. Acronyms for institutions where speci- mens are deposited are: American Muse- um of Natural History, New York (AMNH); Museo de Invertebrados G. B. Fairchild, Universidad de Panama (MIUP); Instituto Nacional de Biodiversi- dad, Costa Rica (INBio). The abbreviations T and S are used for tergum and sternum, respectively. Pappognatha Mickel Pappognatha Mickel 1939: 330-331. Type species: Mutilla pertyi Dalla Torre, 1897, by original designation. Diagnosis. — Pappognatha is the only ge- nus of New World Mutillidae that has males and females with entirely tomen- tose mandibles, the tomentum consisting of short recumbent pubescence, except for the glabrous tips (Mickel 1939). Both sexes have large quadrate heads, broader than the mesosoma, black integument (except T2 maculations in females) and are about equal in body length (11-14 mm). The males have parameres (Figs. 11, 12) straight, broadest at the base, narrowing towards the apex, triangular in lateral view; volsella (Fig. 13) with cuspis stout, broad and somewhat flattened, with rounded apex; digitus short and stout; pe- nis valve (Fig. 14) without preapical tooth. KEY TO MALES OF PAPPOGNATHA MICKEL 1. Clypeus anterior margin broadly nasutiform, with median U-shaped emargination (Figs. 3, 4) 2 -. Clypeus with broad anterior projection, with either shallow concavity or V-shaped emar- gination in the middle of anterior margin (Figs. 1,2) 5 2. Vertex mostly covered with sparse black setae; clypeus middle emargination about Va clyp- eus length 3 -. Vertex mostly covered with sparse pale setae; clypeus middle emargination deeper, nearly % clypeus length 4 3. Clypeus as in Fig. 3; mandible at base with broad, triangular, laminar ventral process (Fig. 5); tibial spurs of mid- and hind legs brown; Central America myrmiciformis (Cameron) -. Clypeus as in Fig. 4; mandible at base with a broad, rounded, laminar ventral process (Fig. 6); tibial spurs of mid- and hind legs pale; Brazil patruelis (Andre) 4. Mesosternum with distinct, slightly transverse tubercle in front of each middle coxa; an- terolateral area of T2 with distinct spot of appressed, pale setae; Brazil carmo (Cresson) -. Mesosternum with distinct, oblique ridge in front of each middle coxa, inner end of ridge almost reaching the margin of coxal cavity; anterolateral areas of T2 with sparse, erect, pale setae; Brazil, Peru obliqua Mickel 5. Clypeus anterior margin with shallow concavity, without V- shaped emargination medially (Fig. 1); mandible at base with broad, triangular, laminar ventral process (as in myr- miciformis, Fig. 5); vertex and T3-4 with mostly black setae; dorsoanterior area of pro- podeum with pale setae, sculpture visible; tibial spurs of mid- and hind legs dark brown to black; Panama panamensis n. sp. Clypeus anterior margin convex, with shallow, V-shaped emargination medially (Fig. 2); mandible at base with reduced laminar ventral process (Fig. 7); vertex and T3-4 mostly covered with dense, pale setae; dorsoanterior area of propodeum covered with very ise pale setae, sculpture not visible; tibial spurs of mid- and hind legs pale; Peru . . peruana n. sp. Volume 14, Number 2, 2005 *i Ui Figs. 1-4. Pappognatha, male clypeus, dorsal view. 1, panamensis. 2, peruana. 3, myrmiciformis. 4, patruelis. Pappognatha panamensis Quintero and Cambra, n. sp. (Figs. 1, 11-14, 16) Pappognatha sp., female: Cambra and Quintero (1992:473); Pappognatha n. sp., female: Yanega (1994). Male. — Integument black. Head: very large, quadrate, slightly broader than long; gena, frons, clypeus and mandible with pale setae; vertex with black setae; clypeus broadly concave on anterior mar- gin (Fig. 1); mandible at base with broad, triangular, laminar ventral process; flagel- lomere I 1.5X as long as flagellomere II; frons and vertex with medium-sized punctures, most spaces between punc- tures usually equal to their diameter; ocel- li small, distance between inner eye mar- gin and lateral ocellus approximately 6X greatest diameter of the latter. Mesosoma: with pale setae, except mesonotum, tegula and anterior half of scutellum with black setae; pronotum, mesonotum and scutel- lum with dense, closely-spaced punctures; propleuron with scattered, small punc- tures; dorsal and ventral areas of meso- pleuron convex, separated by deep groove, elevated areas with medium-sized punctures, anterior and posterior areas practically impunctate; metapleuron im- punctate; propodeum reticulate; scutellum convex; mesosternum with distinct pro- tuberance or tubercle in front of each mid- 193 coxa. Legs: with pale setae, apical tarso- mere produced into short lamellate plate covering claw bases; tibial spurs of mid- and hind legs dark brown to black. Wings (as in Fig. 9) infuscated, except basal and subbasal cells with clear area on basal half; stigma poorly developed; marginal cell large, truncate apically; two submar- ginal cells and traces of a third (as in Fig. 9). Metasoma: with pale setae on: dorsal face of Tl, lateral margin of T2, very small lateral area of T3 and T4, all sterna except S7; black pubescence on: remainder of T2, other terga, and S7; anterior face of Tl gla- brous, impunctate, with conspicuous me- dial tooth on each lateral margin; T2 with medium-sized punctures laterally, some- what smaller at the posterior margin, disk glabrous and with punctures scattered; T3 to T6 and S3 to S6 with dense, small punc- tures interspersed; S2 gibbose anteriorly at the middle, with sparse small punctures. Parameres (Figs. 11, 12) broad at base, nar- rowing toward apex, triangular in lateral view, with dense large setae on basal two- thirds of ventral margin; volsella (Fig. 13) with cuspis stout, broad and somewhat flattened, with rounded apex, covered with setae that are larger toward apex; digitus short, stout; penis valve as in Fig- ure 14. Length: 10.5 mm. Female. — Integument black, except T2 with pair of yellow integumental spots. Head: frons, vertex and gena with black setae, remainder with thin pale pubes- cence and scattered, erect, pale setae; frons and vertex densely, distinctly punctate; gena with distinct, separated punctures. Mesosoma: dorsum with black setae, ex- cept anterior margin of pronotum with pale pubescence, mesonotum with broad, transverse band of pale pubescence inter- rupted medially by very fine black pubes- cence, propodeum with lateral pale-pu- bescent stripe, pleural areas entirely pale pubescent; pronotum and mesonotum with medium-sized, closely-spaced punc- tures; propleuron with medium-sized, separated punctures; meso- and metapleu- 194 Journal of Hymenoptera Research ron almost smooth; side of propodeum with medium-sized, separated punctures. Legs: hind tibia externally with series of three short spines, apical spine at tip of long, conical process; lamellate process of apical tarsomere not emarginate medially; tibial spurs pale. Metasoma: Tl with pale pubescence, except anterior face with me- dian triangular area of black pubescence, extending slightly onto dorsal face; T2 with pair of posteromedian, yellow, al- most circular in tegumental spots (Fig. 15), separated by distance almost equal to their diameter; T2 with black setae, except for sparse, pale pubescence on integumen- tal spots, and broad pale pubescence along lateral margin; T3 to T5 with pale pubescence, interrupted medially by nar- row black line; sterna with pale setae. Length: 10.6-14 mm (n = 12). Holotype: male. — PANAMA: Darien Province: Parque Nacional Darien, Estacion Rancho Frio, Pirre, 80m, 16 Nov 2000-17 Jan 2001, R. Cambra, A. Santos (MIUP), Malaise trap. Paratypes (12 females): Parque Nacional Darien, Pirre Station, 100m, 16 Feb 1989, D. Roubik, 1 9 (MIUP); Parque Nacional Darien, Cana — Cerro Pirre trail, 750m, 10 May 2002, D. Curoe, 19 (MIUP). Colon Province: Donoso, Cuatro Callitas, 21- 26 Jul 2001, P. Gonzalez, 1? (MIUP); Pina, Area Pro- tegida San Lorenzo, 5 Sep. 2000, S. Bermudez, 1 9 (AMNH). Panama Province: 9 km N. El Llano, 29 May 1991, F. Hovore, 1? (MIUP); 2 km E. Altos de Utive, Pacora, 800m, 11 Feb 1998, R. Cambra, 1? (MIUP); Capira, Jordanal, 27-30 Jan 2002, P. Gonza- lez, 1 9 (MIUP). Comarca Kuna Yala: Nusagandi, 16 [an 1991, R. Cambra, 1 9 (MIUP). Code Province: Val- le de Anton, 22 Aug 1991, R. Contreras, 19 (MIUP); El Cope, Division Continental, 900m: 1-2 Sep 1990, D. Quintero, 19 (MIUP); 1 nov 1992, A. Aiello, 19 (MIUP). Chiriqui Province: Fortuna, Quebrada Are- na, 1050m, 8-11 Apr 1999, R. Cambra, A. Santos, 19 (MIUP). Diagnosis. — The male of P. panamensis differs from the other species of the genus by the shallow concavity on the anterior margin of the clypeus, without a medial emargination. The males of P. panamensis md P. myrmiciformis are the only known matha with brown to dark brown s on mid- and hind legs. The fe- '. panamensis is similar to P. pertyi (Dalla Torre) from Brazil but differs in the totally black vertex; the mesonotum with a broad, transverse band of pale pubes- cence, narrowly interrupted medially by a line of black pubescence; and T3-5 with pale pubescence, narrowly interrupted medially by a line of black pubescence. The female of P. pertyi has the vertex with an inconspicuous V-shaped spot of pale pubescence; the mesonotum with a trans- verse band of pale pubescence widely in- terrupted medially by black pubescence; and T3-5 with pale pubescence, widely in- terrupted medially by black pubescence. Pappognatha peruana Quintero and Cambra, n. sp. (Fig. 2, 7, 8) Pappognatha sp.: Quintero and Cambra (1996: 339) Male. — Similar to Pappognatha panamen- sis but differing by: clypeus broadly na- sutiform, anterior margin convex and with minute, shallow,V-shaped emargination medially (Fig. 2); mandible (Fig. 7) at base with reduced laminar ventral process; ver- tex with dense pale pubescence, except area around ocelli with black pubescence; dorsoanterior area of propodeum covered with dense pale pubescence, sculpture not visible; T3 and T4 covered with pale pu- bescence, except small medial area with black pubescence; tibial spurs of middle and hind tibiae pale. Length: 12 mm. Holotype: male. — PERU: Madre de Dios Departa- mento, Reserva Manu, Estacion Pakitza, 9 Mar 1992, D. Quintero (MIUP). Paratype: same data as holotype but 26 Feb 1992, R. Cambra, 1 <5 (MIUP). Diagnosis. — Pappognatha peruana differs from the other species of the genus in hav- ing a shallow V-shaped emargination on the anterior margin of the clypeus, and by the reduced basal ventral laminar process on the mandible. Comments. — The holotype was collected resting on a leaf of a shrub about lm above the ground inside the forest, at about 1600 hours. The paratype, collected Volume 14, Number 2, 2005 195 with an insect net, was flying about 3m above the muddy soil of a narrow forest trail, at noon. This is probably the male of either P, speciosa Mickel or P. limes Mickel. One female of the former was collected in Estacion Pakitza (Quintero and Cambra 1996). The latter species is known from similar habitats in Peru and Ecuador, and might be a junior synonym of P. rotifera (Gerstaecker, 1874), from Colombia. Thus, at present it is best to treat these males as a new species until additional evidence is available. Pappognatha myrmiciformis (Cameron, 1897) (Figs. 3, 5, 9, 10, 15) Male. — Similar to P. panam.en.sis but the clypeus has a deep, rounded emargination medially, the depth of which is one third the length of the clypeus (Fig. 3), and the penis valve in a lateral view (Fig. 15) has a sharp angulation in its anterior margin near the base of the apical tooth. Length: 10-10.5 mm. Material examined. — The specimen from Honduras represents a new country re- cord and the northernmost record for this Neotropical genus. HONDURAS: Atlantida, Lancetilla, Tela, 15 May 1995, R. Cave, 19 (MIUP). COSTA RICA, Limon Province: Sector Cerro Cocori, Finca de E. Rojas, 150m, 26 Mar -24 Apr 1992, F. Quesada, 1 9 (INBio); Cerro Tortuguero, l-120m, Jul 1993, R. Delgado, 19 (MIUP). Heredia Province: Estacion El Ceibo, Parque Nacional Braulio Carrillo, 400-600m, Nov. 1989, R. Aguilar, 1 9 (INBio); Feb 1990, C. Chaves, Id (MIUP); Estacion Magsasay, Parque Nacional Braulio Carrillo, 200m, Mar 1991, A. Fernandez, 19 (INBio). Guana- caste Province: Estacion Pitilla, 700m, 9 km S Sta. Ce- cilia, Nov 1988, 1 9 (INBio); same data, Mar 1995, P. Rios, C. Moraga (Malaise), 19 (INBio); same data, May 1989, I. Gauld, 19 (INBio); Estacion Maritza, 600m, lado O. volcan Orosi, Feb. 1992, F. Araya, 19, 1<5 (INBio). Puntarenas Province: Golfito, Reserva Forestal Golfo Dulce, Estacion Agujas, 250-350m, 17 Mar 2000, A. Azofeifa, 19 (INBio); Golfo Dulce, 24 km Piedras Blancas, 2 Jun-Aug 1989, P. Hanson, 1 9 (British Museum of Natural History'). Alajuela Prov- ince: San Cristobal, 600-620m, 15-28 Mar 1998, F. Quesada, 1 9 (INBio). Guanacaste Province: sotob- osque, W. side volcan Cacao, 1100m, Feb 1989, I. Figs. 5-7. Pappognatha, male mandibles. 5, myrmici- formis. 6, patruelis. 7, peruana. Gauld, 1 9 (MIUP). PANAMA, Code Province: Pen- onome, cabecera Rio Riecito with Quebrada Platanal [8° 52' 44", 80° 16' 38"], UOmsnm, 22 Jim 2001, C. Vega, 19 (MIUP). Darien Province: Parque Nacional Darien, Estacion Rancho Frio, Pirre, 80m, 30 Jul - 8 Aug 2002, A. Santos, R. Miranda, 1 9 (MIUP). Other females from Panama were examined by Cambra ^\nJ Quintero (1992), and Yanega (1994). Distribution. — Honduras, Costa Rica and Panama. Comments on sex association.— Extensive samples show the presence of only one species of Pappognatha in Costa Rica, and the capture of male and female at the same place and time support the associa- tion. Comments. — The male specimen from Guanacaste, Est. Maritza, Feb. 1992, has an 196 Journal of Hymenoptera Research Figs. 8-10. Pappogiwkka, right wings. 8, peruana. 9- 10, myrmiciforrnis. aberrantly shaped second submarginal cell (SM2) on the right wing (Fig. 10). The left wing on the same specimen and the wings of the other male specimen of P. myrmiciforrnis examined (Fig. 9) have typ- ical pentagonal SM2, as do all other male Pavpognatha. Pappognatha patmelis (Andre, 1898) (Figs. 4, 6) Male. — Integument black. Head: very large, quadrate, slightly broader than long; gena, frons, clypeus and mandible with pale setae; vertex with black setae; anterior margin of clypeus broadly nasu- tiform, with median U-shaped emargina- tion (Fig. 4); mandible at base with broad, rounded, laminar ventral process; flagel- lomere I 1.5X as long as flagellomere II; frons and vertex with medium-sized punctures, most spaces between punc- tures usually less than their diameters; small, distance between inner eye • ateral ocellus approximately meter of the latter. Meso- soma: with pale setae, except pronotum, mesonotum, tegula and anterior half of scutellum with black setae; pronotum, me- sonotum and scutellum with dense, close- ly-spaced punctures; propleuron with scattered, small punctures; dorsal and ventral areas of mesopleuron convex, sep- arated by deep groove, elevated areas with medium-sized punctures, anterior and posterior areas practically impuncta- te; metapleuron impunctate; propodeum reticulate; scutellum convex. Legs: with pale setae; apical tarsomeres segment pro- duced into short lamellate plate covering the claw bases; tibial spurs of mid- and hind legs pale. Metasoma: with pale setae on: dorsal face of Tl, lateral margin of T2, very small lateral area of T3 and T4, all sterna except S7; black pubescence on: re- mainder of T2, other terga, and S7; ante- rior face of Tl glabrous, impunctate, with conspicuous medial tooth on each lateral margin; T2 with medium-sized punctures laterally, somewhat smaller along poste- rior margin, disk glabrous and with punc- tures scattered; T3 to T6 and S3 to S6 with dense, small punctures interspersed; S2 gibbose anteriorly at the middle, with sparse small punctures. Parameres broad at base, narrowing toward apex, triangu- lar in lateral view, with dense large setae on basal two-thirds of ventral margin. Length: 11.8 mm. Diagnosis. — P. patmelis closely resem- bles P. panamensis and P. myrmiciforrnis but differs by: the shape of clypeus (Fig. 4); rounded basal laminar process on man- dible (Fig. 6); frons and vertex with dense punctures, most spaces between punc- tures usually less than their diameter; pale calcaria of mid- and hind tibia. Material examined. — BRAZIL: Bahia: Ilheus, Jan 1994, J. Franco, 1 9 (MIUP). Minas Gerais: Caratinga, Fazenda Montes Claros [19° 45' S, 41° 50' W], 30-31 Dec 1994, G. Melo, "Nascida de ninho de Euglossa sp. dentro de orificio em tronco no chao em descompos- icao", 29, IS (MIUP). Distribution. — Brazil. Comments on sex association. -Two fe- Volume 14, Number 2, 2005 197 Figs. 11-15. Pappognatha, male genitalia. 11-14. P. panamensis: 11-12: parameres: 11, dorsal view; 12, lat- eral view; 13, volsella, lateral view; 14, penis valve, lateral view. 15. P. myrmiciformis, penis valve, lateral view. Abbreviations: C, cuspis; P, paramere. males and one male were reared from three cells of one nest of Euglossa sp. The females fit the description of P. patruelis (Andre) given by Mickel (1939) who ex- amined eight females, from Bahia (includ- ing the holotype and a paratype), Sao Pau- lo and Espirito Santo, Brazil. The male of P. patruelis is here described for the first time, based on a single male that died at the teneral stage; its wings were not ex- tended and the mesosternum was not sclerotized and was deformed (concave). Pappognatha speciosa Mickel, 1939 (Fig. 16) Material examined. — The specimens from Colombia and Ecuador represent new country records. COLOMBIA: Amazonas, P.N. Amacavacu, 290m, 30 Feb 1988, 1 9 (AMNH). ECUADOR: Oriente, Li- moncocha (00°24'S, 76°36'W), 21 Jul 1970, C. W. Ret- tenmeyer, 19 (Brothers pers. col.). PERU: 39, Quin- tero and Cambra (1996). Distribution. — Colombia, Ecuador and Peru. Comments. — The female from Ecuador has the integumental spots darkish yel- low, and the pale pubescence on T3-5 not interrupted with black (Brothers, pers. comm.); the type specimen has integu- mental spots ferruginous and pale pubes- cence on T3-5 narrowly interrupted me- dially with black. Mickel (1939) mentioned than P. speciosa is similar to P. egregia, but differs only in the size of the integumental spots of T2 (larger in egregia), and in the outer margin of the spots having pale to- mentose pubescence (glabrous in egregia). In specimens of P. myrmiciformis and P. pa- truelis that we have examined, as well as in many other species in different mutillid genera (e.g., Pseudomethoca and Hoplomu- tilla), the integumental spots on T2 are quite variable in size. Thus, we consider the size of the integumental spots on T2 to be an unreliable character for species identification. Thus, egregia and speciosa might represent but a single species. In ad- dition, Pappognatha torquata Mickel, 1939, might be conspecific with P. speciosa but P. torquata is known only from the holotype from Bolivia (no additional label data). According to Mickel (1939), they differ in having the pronotum pubescence entirely pale tomentose in P. torquata and the pron- otum clothed with sparse, black setae in P. speciosa, and the dense pale pubescence on the vertex extending anteriorly along the inner margin of the eye in P. torquata, but not in P. speciosa. Pappognatha egregia Mickel, 1939 Material examined. — The specimens from Colombia and Ecuador represent new country records. COLOMBIA: Amazonas, Puerto Santander, 160m, 7 Sep 1991, 1 9 (AMNH); Meta, Villavicencio, 25 Jul 1938, 1 9 (MIUP); Putumavo, Santa Rosa, Rio San Mi- 198 Journal of Hymenoptera Research Figs. 16-17. Pappognatha females, habitus. 16. P. panamensis. 17. P. speciosa. guel, 1-30 Oct 1970, Malkin, 1 9 (MIUP). ECUADOR: Napo: Cuyabeno, Apr 1986, 1 9 (MIUP); Cuyabeno, Aug 1983, 400m, 19 (MIUP); El Reventador, May 1986, 19 (MIUP); Limoncocha, 24 Jun 1975, 19 (MIUP). BRAZIL: Amazonas: 60 km N. Manaus, 16 Aug 1991, G. Melo, 19 (MIUP); 1905, 19 (AMNH); Manaus, Reserva Ducke, 3 May 1968, 1 9 (MIUP). Distribution — .Colombia, Ecuador, Bra- zil. Pappognatha limes Mickel, 1939 Material examined. — ECUADOR: Pastaza, Cusuimi, Rio Cusuimi, 150 km SE of Puyo, 15-31 May 1971, B. Malkin, 19 (MIUP); Pastaza, Mera, Apr 2002, 19 (MIUP). PERU: Monson Valley, Tingo Maria, 11 Dec 1954, 19 (AMNH). Distribution. — Ecuador and Peru. Comments. — A female from Colombia (along margins of Rio Calima, Restrepo, Valle, 11 Feb 1984, MIUP) differs from the original description in having the pale pu- bescence on T3-T5, interrupted medially black pubescence, as in typical P. nit the frons is covered with pale pubescence, as in typical P. rotifera (Ger- staecker, 1874). We have identified this specimen as P. rotifera, and conclude that P. limes is probably a junior synonym of P. rotifera, a species previously known from only two specimens from Colombia, the holotype from Bogota and a second specimen from Muzo, Boyaca (Mickel 1939). Pappognatlia obliqua Mickel, 1939 Material examined. — The specimens rep- resent a new country record. PERU: Nauta, Amazonas, Carrasco col., 26 (AMNH, MIUP). Distribution. — Peru and Brazil. Comments. — The two additional males from Peru fit Mickel's description of P. ob- liqua. This might be the male of P. pertyi (Dalla Torre) (Mickel 1939), a species known from Para, Brazil. Volume 14, Number 2, 2005 199 ACKNOWLEDGEMENTS We thank the following curators and their institu- tions for loan or donation of specimens used in this study: James M. Carpenter (AMNH), Gabriel Melo (Universidade Federal do Parana), Paul Hanson (Uni- versity of Costa Rica), Daniel Curoe (Palo Alto, Cal- ifornia, USA), Jesus Ugalde and Alvaro Herrera (IN- Bio) for loan of specimens from the Project Biodiver- sity Resources Developments under the auspices of the World Bank and Global Environmental Facility (GEF). Our field work in Darien National Park, Pan- ama, was supported in part by funds from BIODAR- IEN-ANAM and fund No.04-05-94-01 from the Univ- ersidad de Panama; field work in Manu Reserve, Peru, was supported by funds of the Biological Di- versity in Latin America Project (BIOLAT), Smithson- ian Institution. We thank Denis J. Brothers, University of KwaZulu-Natal, South Africa, for the meticulous review, for providing additional specimen data and the photograph of P. speciosa. Our thanks to Annette Aiello, Smithsonian Tropical Research Institute, and Wojciech J. Pulawski, California Academy of Scienc- es, for their thoughful and critical reviews, and the suggestions made to improve the manuscript. LITERATURE CITED Brothers, D. J., Tschuch, G., and F. Burger. 2000. As- sociations of mutillid wasps (Hymenoptera, Mu- tillidae) with eusocial insects. Insectes Sociaux 47: 201-211. Cambra, R. A. and D. Quintero. 1992. Velvet ants of Panama: distribution and systematics (Hyme- noptera: Mutillidae). Pp. 459-78 in: Quintero A., D. and A. Aiello, eds. Insects of Panama and Me- soamerica: Selected Studies. Oxford University Press, Oxford. Mickel, C. E. 1939. Monograph of a new Neotropical mutillid genus, Pappognatha (Hymenoptera: Mu- tillidae). Annals of the Entomological Society of America 32: 329-343. Quintero, D. and R. A. Cambra. 1996. Contribution a la sistematica de las mutilidas (Hymenoptera) del Peru, en especial las de la Estacion Biologica BIOLAT, Rio Manu, Pakitza. Pp.327-357 in: D. E. Wilson and A. Sandoval, eds. Manu: The Biodi- versity of Southeastern Peru. Smithsonian Institu- tion Press, Washington DC. 679 pp. Roubik, D. W. 1989. Ecology and Natural History of Tropical Bees. Cambridge University Press, Cam- bridge. 514 pp. Wheeler, G. C. 1983. A mutillid mimic of an ant (Hy- menoptera: Mutillidae and Formicidae). Entomo- logical News 94: 143-144. Yanega, D. 1994. Arboreal, ant-mimicking mutillid wasps, Pappognatha; parasites of Neotropical Euglossa (Hymenoptera: Mutillidae and Apidae). Biotropica 26: 465-468. J. HYM. RES. Vol. 14(2), 2005, pp. 200-205 Redescription of Pelicope yuccamica Mason (Hymenoptera: Braconidae: Microgastrinae), With Notes on its Unusual Biology and Relationships J. B. Whitfield, A. A. Valerio, and W. Y Choi Department of Entomology, 320 Morrill Hall, 505 S. Goodwin Ave., University of Illinois, Urbana, IL 61801, USA; jwhitfie@life.uiuc.edu Abstract — A more complete illustrated redescription of the monotypic genus Pelicope (type spe- cies P. yuccamica Mason) is provided to assist in phylogenetic placement of this rare and unusual genus. Misinterpretations of its biology are corrected based on host data published after the orig- inal description— this species appears not to be associated with Yucca as previously reported, but instead only with the incurvariid moth Mesepiola specca Davis, a seed-feeder on Nolina parryi Watson (Ruscaceae). The monotypic genus Pelicope was erect- mechanism. Interestingly, he omitted in ed by Mason in his reclassification of the the description a number of observable world genera of Microgastrinae (Mason features that might have helped place the 1981). He included only the new species genus in his new classification. Although P. yuccamica Mason, a highly distinctive, the type series contained males, no men- polished, xanthic species reported as hav- tion of male genitalic features was made, ing only been collected in a restricted re- and a number of other small features rel- gion of southern California at Yucca flow- evant to comparative study of microgas- ers. Mason characterized the genus as trine genera were also not mentioned, having a rare combination of morpholog- The phylogeny and classification of mi- ical traits within the subfamily Microgas- crogastrine genera has remained contro- trinae — the almost totally xanthic body versial and difficult to resolve (e.g. Mason coloration, the absence of the hind wing 1981, Austin 1990, Walker et al. 1990, Mae- vein 2r-m, and the resemblance of the fore to 1996, Whitfield et al. 2002), in part due wing areolet (Fig. 7) to the fossil genus to the lack of data for a number of genera. Eocardiochiles described from Baltic Amber In this context, a more complete descrip- by Brues (1933), make this genus very pe- tion of Pelicope has been of interest for culiar and (in Mason's view) difficult to some time. Further, a number of other as- classify as closely related to any other ge- pects of P. yuccamica have now come to nus. light some of them scattered enough in Mason's (1981) description was based the literature and collections that it has on six specimens, all taken from the same only recently become possible to integrate locality and month in Riverside County them into a fuller picture of this remark- (Upper Deep Canyon at Horsethief Creek able species. in early June), but collected over a 26 year Biology. — Force (1989), in a study of the period. In addition to a verbal description, biology of the incurvariid moth Mesepiola ie provided simple but accurate line specca Davis, whose larvae feed on the ngs of the head, wings, anterior me- seeds of Nolina parryi (previously in Aga- ^ites and female ovipositor vaceae, now reclassified in the Rusca- Volume 14, Number 2, 2005 201 Figs. 1-2. 1, Lateral habitus photo of Pelicope yuc- camica Mason, female, reared from Mesepiola specca. 2, close-up of ovipositor and sheaths. ceae — butcher's broom family — Judd et al. 2002), reared a series of P. yuccamica, from which we have been able to examine spec- imens. Since extensive studies of yucca/ yucca moth biology and coevolution (re- viewed in Pellmyr 2003) have apparently never yielded Pelicope (O. Pellmyr, person- al communication), it seems clear now that the earlier records of Pelicope as being associated with Yucca were based on mis- identifications of Nolina as Yucca, which it does indeed superficially resemble, al- though the two are now in different plant families. Force (1989) discussed an interesting problem for specialist parasitoids attack- ing M. specca. Nolina parryi does not flower (and thus does not produce seeds) every year, but does so irregularly and synchro- nously in local areas. Mesepiola appears to be able to synchronize with Nolina some- how (although it also has two other less common recorded hosts which might oc- casionally fill in). Either Pelicope has other local hosts besides M. specca (or can find the occasional M. specca on its other host plants), or it too has found some way to synchronize its emergence with the flow- ering years of Nolina. Perhaps the latter strategy might explain why so few Pelicope have ever been collected in an area rich in entomologists. Force (1989) also states that Pelicope are likely to be ovipositing into host eggs, al- though he does not discount that they could be finding very early instar larvae. Oviposition into eggs does not appear to be common in Microgastrinae, but is known to occur in some Cotesia and Diol- cogaster species (Ruberson and Whitfield, 1986). Since Force's observations were in- direct in the sense that he drew conclu- sions based on the timing of host ovipo- sition and Pelicope adult activity rather than on direct Pelicope ovipositions, at this point the question must remain open. More recent studies of Pelicope morpholo- gy.— Maeto (1996), in a comparative study of male genitalia (especially o( volsellar structures) in Microgastrinae, partially de- scribed the male genitalia of Pelicope and presented a photo of a slide mounted gen- ital capsule focusing on the digitus and cuspis structure. There were not enough characters in his study to confidently place the genus phylogenetically, but it did have a unique combination of the five charac- ters polarized by Maeto in his discussion. Whitfield et al. (2002) attempted to syn- thesize the available comparative morpho- logical data for microgastrine genera (in- cluding some of Maeto's data and at- tempting to include the features consid- ered bv Mason) and analyzed these data alone and in combination with DNA se- 202 Journal of Hymenoptera Research quence data from three gene fragments. Specimens of Pelicope suitable for molec- ular study were unavailable at that time, but several morphological characters were coded from the type series for the genus that were neither mentioned by Mason (1981) nor observable from his illustra- tions. The preliminary phylogenies based on both morphological and combined data appeared to confirm Mason's speculation that Pelicope represents an isolated, rela- tively basal lineage, but support for this position is still poor. Below we provide a morphological re- description of P. yuccamica, expanding upon the description of Mason (1981) and the character codings of Whitfield et al. (2002) and providing a series of digital photographs and environmental scanning electron micrographs of a female speci- men from Force's (1989) material. It is hoped that these additional data will im- prove the interpretation of the morpholo- gy of Pelicope for future phylogenetic work. In addition, we hope our clarifica- tion of the biology of Pelicope will stimu- late further field studies of this unusual animal. MATERIALS AND METHODS The principal morphological terminolo- gy used in the species description is that of Huber and Sharkey (1993) as well as of Schuh (1989); except for that of the mor- phology of the propodeum, which is used sensu Townes (1969, Fig. E) and Austin and Dangerfield (1992, Fig. 1). The cutic- ular sculpturing terminology follows that of Harris (1979), while the terminology for the wing venation is a variation of the Comstock-Needham system used by Shar- key and Wharton (1997, Fig. 15). Digital color photographs were taken using a JVC GC-QX5HD digital still cam- era mounted on a Leica MZ12/5 stereo- microscope. ESEM photographs were tak- using a Philips XL30 ESEM-FEG elec- microscope. Pelicope yuccamica Mason (Figs. 1-9) Body color. — Light golden yellowish- brown except for distal 1/5 of hind femur; hind tibia (at least apically) and tarsomer- es dark brownish yellow; ovipositor sheaths and antennae beyond scape very dark brown to essentially black; ventral Vz of head anteriorly whitish. Size. — Body length 3.1-3.5 mm. Fore- wing length 3.1-3.4 mm. Head. — Face broad, very weakly sculp- tured; inner margins of eyes not converg- ing ventrally, sometimes even weakly di- verging. Clypeus broad, shallowly semi- circular above broad subtriangular labrum which largely conceals mandibles in fron- tal view. Antennae unusually thin and short for Microgastrinae (Fig. 1); placodes two-ranked on at least proximal 10 flagel- lomeres, and ventral area same as dorsal for subapical flagellomeres. Glossa trun- cate. Maxillary and labial palps slender, pale yellowish, relatively short. Mesosoma. — Pronotal lateral area with two nitid grooves; propleuron lower outer corner simple (Fig. 5). Mesonotum shiny, nearly smooth, without or with very in- distinct puncation, but with very smooth, broad, shallow suggestions of notauli and raised medial lobe (Fig. 4). Epicnemial ca- rinae absent; mesopleura weakly sculp- tured except anterodorsally and near poorly defined sternaulus (Fig. 5). Scutos- cutellar groove narrow, convex towards anterior, with very tiny shallow pits (Fig. 4). Scutellar disc subtriangular, very weakly punctate, slightly wider anteriorly than long; lunulae slightly curved dorsally and mainly of uniform width, with nitid medio posterior scutellar band. Metano- tum with sub-scutellar hairs absent; ante- rior margin appressed to scutellum. Pro- podeum smooth, with more horizontal an- terodorsal area greatly shortened and gradually curving to strongly declivous face, without carinae. Legs. — Fore telotarsus normal in shape Volume 14, Number 2, 2005 203 3H ■ jb* v jJa. Figs. 3-6. ESEM Photos of P. yuccamica female: 3, head, anterior view; 4, mesosoma, dorsal view; 5, meso- soma, lateral view; 6, anterior metasomal tergites, dorsal view. (not excavated apicoventrally). Hind cox- ae not strongly enlarged (thus resembling non-Diolcogaster Microgastrinae); hind tib- ial spurs subequal in length and about xh length of hind basitarsi; tarsal claws ap- pearing simple under low magnification but actually bearing a cluster of 4-5 tiny slender teeth basally (Fig. 9). Wings (Fig. 7). — Forewing vein 3Rs strongly convex towards costal vein, lM + IRs straight, r-m meeting 3Rsa to close the unusually large areolet, 2A ab- sent. Stigma pale yellowish basally, oth- erwise brown; veins mostly yellowish brown. Hindwing veins r, 2A and 2r-m absent, cells 1R and 2R of approximately same width, veins 2Cu and cu-a straight; vannal lobe with a distinct apical notch, convex in shape and with setosity short and sparse beyond broadest point. Metasoma. — First metasomal tergite with gentle anterior basal excavation rather than mediobasal longitudinal groove (Fig. 6), apical half flat to gently arched and without medioapical excavation, in dorsal view narrowing towards apex of tergum. Second metasomal tergite subtriangular and relatively small, appearing undefined posteriorly due to exceptionally weak pos- terior groove. Third metasomal tergum 1.5X as long as second tergite. General as- pect of metasoma not carapacelike and with terga nitid. Medial longitudinal area of hypopygium desclerotized and with longitudinal pleats. Ovipositor mecha- nism (Fig. 8) with eighth tergite 3x taller (dorsoventrallv) than long and with the apodeme shorter than wide; second val- vifer tall and lorate; the slender straight ovipositor sheaths (Figs. 2, 8) arising from second valvifer near apex, setosity present through length; ovipositor in lateral view very weakly decurved and evenly tapered throughout its length. Male genitalia (from Maeto, 1996 — his Table 1 and Fig. 10). — Lamina volsellaris 204 Journal of Hymenoptera Research Figs. 7-9. P. yuccamica, female: 7, wings; 8, ovipos- itor mechanism, lateral view; 9, hind tarsal claw. 7, 8 after Mason (1981); used with permission. 0.28 mm long, with 2 setae. Digitus 0.57 X as long as lamina volsellaris, slender ba- sally and apically acute and directly dor- sally, but with ventral edge slightly con- vex; apex with 3-4 apical teeth. Larva and cocoons. — Force (1989) states that Pelicope spend the winter in the soil in the host larvae and adults emerge the following late May/early June when the adult Mesepiola are ovipositing into devel- oping seeds. Since the host spins a tough cocoon in the soil, probably the parasitoid cocoon is inconspicuous within it. Material examined. — Holotype: USA, California, Riverside Co., Upper Deep Canyon at Horsethief Creek, 1000 m, ll/vi/1965, Col. E. Schlinger. Para- types (2 ? 9 and 3 6 6) and holotype in the U. S. National Museum. Additional specimen used for re- discription and figures: USA: California: Riverside Pinyon Flat, 18/vi/1987, D. Force, ex. Mesepiola Una parryi (in Illinois Natural History ACKNOWLEDGMENTS We would like to thank Drs. Henri Goulet (Cana- dian National Collection, Ottawa) and David Smith (U. S. National Museum, Washington) for the oppor- tunity to examine Pelicope specimens in their care. Olle Pelmyr provided invaluable help on the question of whether Pelicope is associated with Yucca moths, and helped locate the Force paper. Scott Robinson of the Imaging Technology Group, Beckman Institute, University of Illinois, provided assistance with the ESEM photography. This work was supported in part by National Science Foundation grant DEB 0235784 and U. S. Department of Agriculture NRI grant 2003- 35316-13679 to JBW. LITERATURE CITED Austin, A. D. 1990. Revision of the enigmatic Austra- lasian genus Miropotes Nixon (Hymenoptera: Braconidae), with comments on the phylogenetic importance of the female ovipositor system. Sys- tematic Entomology 15: 43-68. Austin, A. D. and P. C. Dangerfield. 1992. Synopsis of the Australasian Microgastrinae (Hymenop- tera: Braconidae), with key to genera and de- scription of new taxa. Invertebrate Taxonomy 6: 1- 76. Brues, C. T. 1933. The parasitic Hymenoptera of the Baltic amber. Berstein Forschungen 3: 4-178. Force, D. C. 1989. Observations on the parasitoids of Mesepiola specca Davis (Lepidoptera: Incurvari- idae). Pan-Pacific Entomologist 65: 436^439. Harris, R. A. 1979. A glossary of surface sculpturing. Occasional Papers of the Bureau of Entomology of the California Department of Agriculture. No. 28, 32 pp. Huber, J. T. and M. J. Sharkey. 1993. Structure, Chap- ter 3. Pp. 13-59 in: Goulet, H. and Huber, J.T., eds. Hymenoptera of the World: An Identification Guide to Families. Agriculture Canada Publication 18894/E, Ottawa. judd, W. S., C. S. Campbell, E. A. Kellogg, P. F. Ste- vens, and M. J. Donoghue. 2002. Plant Systemat- ica: A Phylogenetic Approach. Second Edition. Sin- auer Associates, Sunderland, MA. Maeto, K. 1996. Inter-generic variation in the external male genitalia of the subfamily Microgastrinae (Hymenoptera, Braconidae), with a reassessment of Mason's tribal system. Journal of Hymenoptera Research 5: 38-52. Mason, W. R. M. 1981. The polyphyletic nature of Apanteles Foerster (Hymenoptera: Braconidae): a phylogeny and reclassification of Microgastrinae. Memoirs of the Entomological Society of Canada, No. 115, 147 pp. Pellmyr, O. 2003. Yuccas, yucca moths and revolu- tion: a review. Annals of the Missouri Botanical Garden 90: 35-55. Ruberson, J. R. and J. B. Whitfield. 1996. Facultative Volume 14, Number 2, 2005 205 egg-larval parasitism of the beet armyworm, Spodoptera exigua (Lepdioptera: Noctuidae) by Cotesia marginiventris (Hymenoptera: Braconi- dae). Florida Entomologist 79: 296-302. Schuh, R. T. (Ed.) 1989. The Torre-Bueno Glossary of Entomology (revised edition). The New York En- tomological Society, New York, 849 pp. Walker, A. EC, I. J. Kitching, and A. D. Austin. 1990. A reassessment of the phylogenetic relationships within the Microgastrinae (Hymenoptera: Bra- conidae). Cladistics 6: 296-306. Sharkey, M. j. and R. A. Wharton. 1997. Morphology and terminology. Pp. 19-37 in: Wharton, R. A, P. M. Marsh, and M. J. Sharkev, Eds. Manual of the New World Genera of the Family Braeonidae (Hy- menoptera). Special Publication of the Internation- al Society of Hymenopterists, No. 1 . Whitfield, J. B., P. Mardulyn, A. D. Austin, and M. Dowton. 2002. Phylogenetic relationships ami >ng microgastrine braconid wasp genera based on data from the 16S, COI and 28S genes and mor- phology. Systematic Entomology 27: 337-359. J. HYM. RES. Vol. 14(2), 2005, pp. 206-207 NOTE Ormyrns salmanticus Nieves-Aldrey, 1984 (Hymenoptera: Chalcidoidea: Ormyridae), a Valid Species J. L. Nieves-Aldrey and M. Hernandez Nieves Museo Nacional de Ciencias Naturales, Departamento de Biodiversidad y Biologfa Evolutiva, Jose Gutierrez Abascal 2, 28006 Madrid, Spain; email: aldrey@mncn.csic.es Ormyrus salmanticus Nieves-Aldrey was described in 1984 from Casillas de Flores, Salamanca (Spain), reared from galls of Aulacidea subterminalis Niblett, 1946, a herb gall wasp (Hymenoptera, Cynipidae, Ay- lacini) that induces semi-subterranean galls in runners of Hieracium pilosella L. (Asteraceae). This species is easily distin- guished from other Iberian ormyrid spe- cies by several important diagnostic char- acters, as follows: second metasomal ter- gite smooth dorsally and with weak cren- ulate sculpture of tergites 3-7, lacking the typical rows of deep foveae of many Or- myridae; head only slightly broader than high in anterior and posterior views; post- marginal vein relatively long, 1.6 times as long as stigmal vein; metatibia as long as metatarsus. Narendran (1999) published a system- atic monograph on the Indo-Australian Ormyridae and stated that O. salmanticus is a junior synonym of Ormyrus langlandi Girault, 1920, a decision that has been sub- sequently accepted by other authors (Noyes 2002). Narendran did not examine the type material of O. salmanticus that is available at the Museo Nacional de Cien- cias Naturales in Madrid. However, the author said that "from the description and figures of Ormyrus salamanticus Aldrey (Aldrey, 1984) (sic), described from Spain, is evident that it is a new junior syno- i of Ormyrus langlandi Girault". spite its superficial resemblance, on the basis of the original description and figures of O. salmanticus, there are how- ever many morphological features of this species that do not fit the description giv- en by Narendran of Ormyrus langlandi. The specific biology of O. salmanticus and its more restricted geographical distribu- tion compared with O. langlandi (the for- mer recorded only from Spain, although probably much widely distributed in Eu- rope, and the host gall and plant distri- bution do not overlap the range of O. lang- landi), were also additional data that cast doubts on the validity of this synonymi- zation. To demonstrate it, we borrowed 16 specimens of O. langlandi housed at the NHM of London and compared them with O. salmanticus. Examined specimens came from Zim- babwe (former Rhodesia) labelled as O. de- caryi Risbec, 1955, a species synonymized with O. langlandi by Boucek (1988). We tried to examine first Girault's O. langlandi material from the QLD museum but un- fortunately we did not receive any replies to our letters. Comparison of series of specimens of the two supposedly conspecific species showed important morphological differ- ences between them and thus clearly in- dicated their separate specific status. Considering the evidence provided above we propose the re-establishment of O. salmanticus Nieves-Aldrey as a valid species, stat. res. Volume 14, Number 2, 2005 207 Table 1. Major morphological differences between O. langlandi and O. salmanticus. Diagnostic characters O. salmanticus 1 male, 3 Females examined O. langlandi 6 males, 10 females examined Female antenna Male antenna Forewing Metasoma Biology and distribution Two anelli but the second anellus bears a single placodeal sensilla and is 1.3 times as long as wide; flagellar seg- ments 2 to 5 clearly transverse, about 1.4 times as wide as long. Placodeal sensillae of flagellar segments ar- ranged in a single row. Flagellar segments transverse. Placo- deal sensillae arranged in a single row. Postmarginal vein relatively longer; marginal vein 1.2 times as long as postmarginal; postmarginal vein 3.2 times as long as stigmal vein. Sculpture on dorsal part of second me- tasomal tergite of female absent; ter- gite completely smooth. A parasitoid of Aulacidea subterminalis Niblett that induces galls in runners of Hieracium pilosella L. Recorded only from Salamanca and Zaragoza (Spain) but probably much more widely distributed following its host gall wasp and plant. Two anelli; second anellus transverse, about 2 times as wide as long; flagel- lar segments 2 to 5 clearly longer than wide, about 1.3 times as long as wide. Placodeal sensillae of flagellar segments arranged in two rows. Flagellar segments longer than wide. Placodeal sensillae smaller, arranged in four or five rows. Postmarginal vein relatively much shorter; marginal vein 3.5 times as long as postmarginal; postmarginal vein 1.6 times as long as stigmal vein. Dorsal part of second metasomal ter- gite of female with strong reticulate sculpture. Biology unknown. Recorded from Aus- tralia, India, Laos, China, Zimbabwe and Madagascar. LITERATURE CITED Boucek, Z. 1988. Australasian Chalcidoidea (Hyme- noptera). A biosystematic revision of genera of fourteen families, with a reclassification of species. CAB International, Wallingford, Oxon, U.K. 832pp. Narendran, T. C. 1999. Indo-Australian Ormyridae (Hy- menoptera: Chalcidoidea). Privately published, T.C. Narendran, Department of Zoology, University of Calicut, Kerala, India. iii+227pp. Nieves-Aldrey, J. L. 1984. Primeros datos sobre los representantes de la familia Ormyridae en 1 ^~ pana, con description de una nueva especie (Hym., Chalcidoidea). Graellsia 40: 119-127. Noyes, J. S. 2002. Interactive Catalogue of World Chal- cidoidea (2001 — second edition). CD-Rom. Taxapad and The Natural History Museum, London. INSTRUCTIONS FOR AUTHORS General Policy. 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