Trigonalidae (Hymenoptera) of Thailand, other southeastern Asian records, and a new Trigonalys from India

Seven species of Trigonalidae are recorded from Thailand, Lycogaster flavonigrata Chen, Achterberg, He & Xu, L. rufiventris (Magretti), Taeniogonalos fasciata (Strand), T. gestroi (Schulz), T. sauteri Bischoff, T. tricolor Chen, and an unidentified species. Four species, L. flavonigrata, T. cordada Chen, Achterberg, He & Xu, T. sauteri, and T. tricolor are newly recorded from Laos. Diagnoses, illustrations, and distribution records are given for the Thailand species. Trigonalys india Smith, sp. n., is described from southern India for specimens previously misidentified as T. rufiventris.


Introduction
Trigonalidae is a small, worldwide family with the greatest diversity in the tropical regions. However, there are few records from southeastern Asia, most of which are recorded in Carmean and Kimsey (1998) and Chen et al. (2014). Previously, only two species were known from Thailand, T. gestroi (Schulz, 1908) (as Taeniogonalos thwaitesii (Westwood, 1874)) and T. tricolor Chen, 1949, and only one species from Laos, T. gestroi. Here, with lateral longitudinal white stripe; tergite 3 with lateral oval white spot; tergite 4 with small lateral white spot, much smaller than that on tergite 3; sternite 2 with apical yellow transverse narrow band. Wings darkly infuscate; lighter toward base; veins and stigma black. Antenna short, flagellum slightly incrassated at center. Medio-apical process present on sternite 2, slightly concave at center.
Distribution. This species was described from China (Fujian, Jiangxi, Yunnan) (Chen et al. 2014). These are the first records from Laos and Thailand.
Comments. The almost completely black color with some yellow markings on the metasoma and the dark wings help distinguish this species.
Lycogaster are fairly rare in collections. From observations by PT, a good number of species of Asian Lycogaster seem to parasitize the Eumeninae through caterpillars, somewhat like Bareogonalos. At least in Korea this explains their scarcity in collections. They are almost never found in traps set in forest but seem to research more sunny habitats where potter wasps nest, and therefore are likely to have a much better resistance to desiccation than the other Trigonalids.  Trigonalys rufiventris Magretti 1897: 311, figs 2, 3; Carmean and Kimsey 1998: 70. Lycogaster rufiventris : Schulz 1908: 28, fig. 2;Bischoff 1938: 10;Weinstein and Austin 1991: 428;Chen et al. 2014: 45. Diagnosis. Female: Length, 8.0-10.0 mm. Antenna black. Head black with clypeus except for black narrow anterior margin and mandible except black apex white. Mesosoma black with anterior margin of pronotum, spot on posterior lateral corners (near tegula) of pronotum, tegula, small oval spot on posterior-central prescutum, oval spot on anterocentral area of mesoscutellum, and metascutellum white. Legs black with most of trochanters except ventrally, extreme tip at base and apex of femora, outer surface of fore tibia, foretarsus, and extreme bases of mid-and hind tibiae white. Metasoma orange with tergite 5 or 6 to apex, and sterna 3 to apex black; apical broad margin of sternite 2 white. Wings lightly infuscated, a little darker toward apex; veins and stigma black. Head: Antenna with 24-25 antennomeres; flagellum somewhat incrassate at center. Shiny; vertex almost without punctures; frons more densely punctate; covered with fine white hairs. Mesosoma: Shiny, uniformly densely punctate; covered with fine white hairs. Metasoma: Shiny, uniformly densely punctate; covered with fine white hairs. Medio-apical process with small indentation at center on sternite 2 (Figs 4,6).
Comments. This was the most commonly collected species and can be distinguished by its yellow and black color pattern and the female lacking a medio-apical process on the second sternite. Chen et al. (2014) discussed the use of the name gestroi. This was commonly known as T. thwaitesii (Westwood), which was described from Sri Lanka and which Carmean and Kimsey (1998) believed to be a widespread, variable species in Asia. Examination of Westwood's type (OXUM) indicated that thwaitesii is different from specimens from southeastern Asia which were previously referred to as T. thwaitesii, confirming Schultz's (1908) conclusions. The name gestroi is thus available for the Asian populations. So far as we can tell, T. thwaitesii is confined to Sri Lanka and possibly southern India. Further studies are needed based on specimens from its entire range.  Taeniogonalos sauteri Bischoff 1913: 153. Diagnosis. Female, length, 8.0 mm. Black with yellow pattern similar to Figs 14-16. Head with yellow markings on occiput, gena, inner orbits, antennal tubercles, and clypeus. Mesosoma with yellow spots at anterior corners of middle lobe and spot on upper pronotum; meso and metascutellum orange. Legs black with hind coxa and bases of tibiae white. Metasoma black with broad apical yellow bands on first and second segments and lateral yellow spots on third to apical tergites. Short, blunt medio-apical process, truncate at apex, on sternite 2 (Figs 14,16). Male similar to female except for sexual differences.
Comments. These specimens are identical to T. sauteri as characterized by Chen et al. (2014). Distribution. China, Thailand, Korea (Chen et al. 2014). Recorded from Thailand by Carmean and Kimsey (1998) and Chen et al. (2014), but no localities given. Chen et al. (2013) recorded 11 provinces in China but not Sichuan.
Comments. This is similar to T. sauteri but the metasoma is almost completely black and there is a distinct medio-apical process on the second sternite. This process is short and apically truncate. It does not key in Chen et al. (2014) and we do not have enough specimens to evaluate its status. Fig. 23 Specimen examined. THAILAND Nan, Doi Phu Kha NP Office 6, 19°12.349'N, 101°4.617'E, 1360m, Malaise trap, 22-19.iii.2008.

Taeniogonalos sp.2
Comments. This may be male of the unidentified female above. The antenna has elongate tyloids on antennomeres 10-15.

Description.
Female. Length 8.0 mm. Head black; clypeus and mandibles yellowish except apex of mandible black. Mesosoma black with yellow spot on dorsoposerior pronotum. Legs black with following yellow: apices of coxae, trochanters, extreme bases of femora, apical third of fore femur, outer surfaces of tibiae. Metasoma orange with first segment, and central area of sterna 2 and 3 black. Basal half of wings hyaline, somewhat darkly infuscated at center and apex; veins and stigma black. Head and body with silvery hairs. Head smooth, shiny with very few scattered fine punctures; mesosoma and metasoma uniformly finely punctate. Antenna long, filiform, with 25-26 antennomeres, 2× head width. Malar space narrow, slightly less than diameter of an ocellus. Eyes small, round, inner margins not converging below; lower interocular distance about 1.4× eye height. Lower frons between antennae shelflike or ledgelike, overhanging supraclypeal area. Distance between toruli about equal to distance be-tween torulus and inner margin of eye. Occipital carina narrow, less than half diameter of an ocellus. Medio-apical process at apex of sternite 3, flangelike, directed backward apically, quadrate.
Etymology. Named after the country of origin, a noun in apposition. Comments. These are the specimens examined and identified as Trigonalys rufiventris (Magretti) by Carmean and Kimsey (1998). Generic characters place them in Trigonalys, and, although similar to L. rufiventris in color, they differ by the placement of the occipital carina, overhanging shelflike interantennal area, presence of tyloids on the antenna of the male, and a medio-apical process on the third sternite of the female.
Most species of Trigonalys are Neotropical and Ethiopian. Only one species is recorded from Asia, T. lachrynosa Westwood, 1874, described from Mindanao, Philippines. We have examined one female of T. lachrynosa from "Surigao, Mindanao" in the USNM, identical to Westwood's illustration (1874, plate 23, fig. 5), the same specimen examined by Carmean and Kimsey (1998). Trigonalys lachrynosa is almost entirely black with small yellow spots on the mesosoma. We know of no other species of Trigonalys with the color combination of T. india -the mostly black head, mesosoma, and legs and a mostly reddish metasoma.    Bradley (1917) to be closely related as they appeared next to each other in his key. Malloch (1926) grouped these same species (aequalis Fox, 1899;idiotes Cockerell, 1895;noctivaga Bradley,1917, andnormalis Malloch, 1926) in what he called the aequalis group of Brachycistis. Wasbauer (1958) in his preliminary studies of the subfamily Brachycistidinae discovered a new species (lamellatus Wasbauer, 1958) from Baja California, Mexico, which because of its unique combination of traits was set aside as representing a possible new genus. When this species was compared to others in the genus Brachycistis, the species Malloch had placed in the aequalis group were found to share these same traits. To accommodate these species, Wasbauer proposed the genus Acanthetropis and included it in his key to the genera known at that time (Wasbauer 1966). The most recent key to genera can be found in Kimsey and Wasbauer (2006).

Materials and methods
Type repositories and specimens studied below are indicated by the following acronyms: ANSP -Academy of Natural Sciences of Philadelphia, Philadelphia, Pennsylvania, USA; BME -Bohart Museum of Entomology Morphological terminology follows that of Wasbauer (1966). Of the head, LID = lower interocular distance, the shortest distance between the lower margins of the compound eyes. UID = upper interocular distance, the shortest distance between the upper margins of the compound eyes. The abbreviations L, W and H refer to length, width and height respectively. Diagnosis. Species of Acanthetropis are large, relatively heavy-bodied brachycistidines. The males can be immediately distinguished by the presence of a scrobal sulcus, very long marginal cell of the forewing and metasomal sternum II with a median longitudinal carina or fold. In addition the frons bears a carina above and laterad of the antennal sockets. Male description. Body length 6.5-17.0 mm. Head. Rounded, broader than long; ocelli enlarged with a transverse sulcus between their posterior margins; compound eyes slightly convergent below, inner margins broadly emarginate; antennal socket rim with broad carina or thickening beneath small carina above and lateral to antennal socket; clypeus transverse, not projecting apically, clypeal L/W ratio not exceeding 0.4, central part slightly convex, with narrow, apically projecting ledge; mandibles tridentate, weakly developed external carina extending from mandibular base to point just proximad of inner tooth, obsolescent near its middle; maxillary and labial palpi welldeveloped, conspicuous, 6 and 4 segmented respectively. Mesosoma. Robust, moderately long; pronotum narrow, transverse, nearly vertical, humeral angles rounded, not prominent; mesonotum with parapsidal furrows long, strongly impressed; scutellum with or without median longitudinal sulcus; mesepisternum carinately produced or with low, rounded protuberance anterodorsally, irregularly convex; propodeum with dorsomedian longitudinal sulcus, area laterad of sulcus sloping abruptly posteriorly, dorsal and posterior propodeal faces separated by transverse carina; coxae simple, not carinate ventrally, with well-developed admesal stridulatory area; wings long, venation well-developed, forewing with three submarginal and two discoidal cells, third submarginal cell two-thirds or more length of second, first transverse cubital vein arising beyond basal third of first submarginal cell, marginal cell elongate, costa extending distad of stigma a distance greater than half length of stigma; hindwing with radial and cubital veins well-developed, cubitus arcuate, forming angle with transverse cubital of less than 135°, jugal lobe much shorter than submedian cell. Metasoma. Segment I narrower than II in dorsal view, sternum I with median sulcus somewhat expanded posteriorly; sternum II with strong basal median longitudinal carina or fold. Genital capsule. Paramere broadest subapically, ventrally rounded to acute apex; aedeagus slender, linear or gradually expanded toward apex; volsella with cuspus broadly rounded, digitus produced apically into short, blunt point in ventral view, inner margin of volsella with several stout spines and slender setae. Color. Reddish brown to dark brown.
Female. Unknown, although the female of Acanthetropis may well be the "Genus A" described by Kimsey (2005).
Distribution . Acanthetropis is known from geographically scattered records in the USA: Arizona, California, Idaho, Kansas, Nebraska, Nevada, New Mexico, South Dakota, Utah, and in Mexico: Baja California, Baja California Sur, Sonora, Hidalgo and Zacatecas. Further collecting in intervening areas will probably reveal a more widespread fauna. Second metasomal sternum with a median longitudinal elevation, terminating in a spine (Figs 5,10); gular carina sharply elevated anteriorly, forming a tooth visible behind the posterior mandibular condyle in full lateral view (Figs 3,4 Flagellomere I at least 3.5× as long as greatest width (Fig. 8); metasomal segment I long and petiolate, L/H 2.5 or more ( Fig. 9) ....... idiotes (Cockerell) -Flagellomere I less than 3.0× as long as greatest width (Fig. 7) Diagnosis. This species most closely resembles A. idiotes. It can be distinguished by the shorter antennomeres, mesepisternum without vertical or transverse ridge, the shorter first metasomal segment and sharp median longitudinal ridge of metasomal sternum I. Male description. Body length 6.5-13.0 mm. Head. Flagellum with appressed yellow pubescence; L/W ratio of flagellomere I 2.4-2.9 (Fig. 7); lower rim of antennal socket produced into distinct vertical carina beneath; clypeus with scattered punctures, becoming larger and irregular apically; gular carina elevated for most of length but not produced into an anterior tooth. Mesosoma. Pronotum with small punctures medially, larger, coarser laterally; mesepisternum with a low, rounded prominence anteriorly, not carinately produced, mesonotum with small, scattered punctures; propodeum with dorsum finely reticulate, impunctate, dorsal sulcus with edges scarcely or not appearing carinate, posterior transverse sulcus absent or poorly defined, without or with anterior ill-defined carina; posterior transverse carina often incomplete. Metasoma. Segment I short and stout, L/H 1.3-1.5 sternum with longitudinal sulcus expanded and deepened posteriorly, tergum heavily punctate; tergum II moderately to lightly punctate, punctures small and shallow; terga III-VII with punctures small, shallow except for row of larger, deeper punctures before apex; sternum II with median basal longitudinal ridge terminating in blunt spine (as in Fig. 6). Genital capsule (Fig. 11). Color. Uniform medium brown. Entire body clothed with long, golden setae.

Key to the species of Acanthetropis
Distribution ( Seasonal distribution. This is a summer species, collected from June through September. Discussion. When Malloch proposed A. normalis as a new species in Brachycistis, he had only seen one specimen of Brachycistis aequalis Fox and thus was unaware of the amount of structural variation seen in that species. The differences he perceived in the dimensions of the first metasomal segment proved to fall within the normal range of variation of this trait in A. aequalis. Figs 8,9,12,16 Brachycistis idiotes Cockerell 1895a: 63. Holotype male; USA: New Mexico (Repository unknown).

Diagnosis.
Acanthetropis idiotes is separable from other species in the genus by the rounded, unridged mesepisternum, lack of a gular elevation, long antennae, and long, slender body. Male Description. Body length 12-17 mm. Head, with few short to moderately long setae, clypeus and mandibles with more numerous, longer setae; scape with shorter setae; flagellum with tiny, scattered setae, flagellomere I L/W 3.9-4.2 (Fig. 8); lower rim of antennal socket with low carina beneath; clypeus with uniformly scattered punctures; gular carina slightly elevated but not produced into tooth anteriorly. Mesosoma. Pronotum with scattered, small punctures medially, larger, denser punctures laterally; mesepisternum with large, scattered punctures, becoming nearly impunctate posteriorly, with low, rounded prominence anteriorly, not carinately produced; mesonotum and scutellum with large, scattered punctures; propodeal dorsum impunctate, dorsal sulcus not usually carina margined, areas laterad of sulcus poorly defined, posterior transverse sulcus absent or ill-defined without anterior carina, posterior transverse carina complete. Metasoma. First segment long and slender, L/H 2.5-4.0 ( Fig.  10), sternum with longitudinal sulcus narrow, slightly widened posteriorly; tergum I slightly rugose, with small, irregular punctures laterally; tergum I with small, scattered punctures; terga III-VII nearly impunctate, except for row of shallow, setose punctures apically; sternum II with a median basal longitudinal ridge terminating in blunt spine (as in fig. 9). Genital capsule (Fig. 12). Color. Uniform medium to chestnut brown, entire body shining, clothed with long golden setae. Seasonal distribution. Specimens of this species have been collected only during the winter months (November through January).
Discussion. Collection of desert insects during the winter in temperate areas of the western U.S. has been largely ignored because of assumptions about seasonality. However, a unique fauna of brachycistidine wasps has developed with adult males appearing during the winter. In addition to Acanthetropis idiotes, two species of Brachycistis, juncea Wasbauer, 1966, andlongula Wasbauer, 1966, in the petiolata group are known only from winter collection dates (November and January) (Wasbauer 1966). Interestingly, the habitus of males of these two species is similar to that of A. idiotes males. They are longer and more slender than males of other species in their respective groups. Figs 1,6,13,17 Acanthetropis lamellatus Wasbauer 1958: 140. Holotype male; Mexico: Baja California Sur, La Paz (CAS).

Diagnosis.
The most distinctive feature of lamellatus is the elevated transverse ledge on metasomal sternum II. Additional diagnostic features include the high sharp mesepisternal prominence and gular carina not elevated or dentate. Male description. Body length 11-14 mm. Head. Face shining with few scattered punctures, punctures stronger between lateral ocelli and on occiput; scape with few long, erect setae ventrally; flagellomere I L/W 2.5-3.1; lower rim of antennal socket not sharp or carinate below; basal half of clypeus shining, impunctate, apical half with several irregularly spaced, broad, shallow punctures and long apically directed setae; gular carina not abruptly raised anteriorly, not visible below mandibular condyle in full lateral view; mandibles with several long, stout amber-colored setae on outer surface. Mesosoma. Strongly punctured, sparsely clothed with medium to long erect or suberect setae; anterior face of pronotum shining medially, nearly impunctae, lateral surface strongly punctate; mesonotum with punctures smaller, more closely set anteriorly; mesepisternum with strongly raised, ridge-like elevation below anterodorsal protuberance; propodeum with dorsal sulcus strongly impressed, shining, raised area laterad of sulcus finely reticulate, impunctate except for small lateral area, posterior declivity sharp, carinate, sulcus between declivity and posterior transverse carina smooth, shining; posterior propodeal face separated from lateral face by lateral diagonal carina meeting posterior carina dorsally, obsolete before posterior rim. Metasoma. Segment I short, stout, L/W 1.7-2.3 in lateral view; sternum II with basal longitudinal fold, terminating at raised, transverse lamella (Fig. 6). Genital capsule (Fig. 13). Aedeagus widest near subapically, narrowed to bluntly pointed apex. Color. Uniform medium to chestnut brown, entire body shining, clothed with long golden setae; flagellum sparsely clothed with minute appressed whitish pubescence.  (Figs 3,4), with relatively sparse, short setae, mainly on clypeus and outer surface of mandibles, longest on occiput, scape and inter-antennal area with moderately long setae; clypeus with large, irregular punctures, flagellomere I L/W 3.0-3.5; lower rim of antennal socket with broad carina or swelling beneath; clypeus with large, irregular punctures; gular carina produced into prominent tooth visible below posterior mandibular condyle. Mesosoma. Pronotum anterior face with very small punctures, humeral angles and lateral face with large, closely set, irregular punctures; mesepisternum heavily punctate, punctures large, close set, irregular; mesepisternal prominence high and sharp, appearing carinate in certain angles; mesepisternal carina present; mesonotum and scutellum with scattered punctures, scutellum usually with median longitudinal sulcus extending from anterior to posterior margins; propodeal dorsum impunctate, dorsal longitudinal sulcus carinately margined, areas laterad of sulcus well-defined, margins usually appearing carinate, posterior transverse sulcus impunctate, often traversed by longitudinal ridges, posterior transverse carina well-developed, complete. Metasoma (Figs 5,10). Segment I short and stout, L/H 1.5-1.9, sternum I with longitudinal sulcus gradually expanded and deepened posteriorly; tergum I heavily punctate, punctures large, often irregular; terga II-VII with scattered large and small punctures and row of setose punctures before apex of each segment; sternum II with median basal longitudinal ridge terminating in blunt spine (Fig. 5). Genital capsule (Fig. 14). Color. Uniform medium to dark brown, body shining, heavily punctate, entirely clothed with shiny white to straw-colored setae; flagellum with tiny erect to appressed golden setae. Distribution (Fig. 18 Geographic distribution of Labidus coecus (latr.) (Hymenoptera, Formicidae), a subterranean army ant James K. Wetterer 1 , Gordon C. Snelling 1

Introduction
Labidus coecus (Latreille) (Hymenoptera, Formicidae) is a widespread New World subterranean army ant. Longino (2007) wrote that L. coecus: "is one of the most remarkable of all army ant species. It has an extremely broad ecological tolerance. It occurs across a great latitudinal range, from the equator to the subtropics of both North and South America. It occurs in dry forest and wet forest, in primary forest and in second growth, in coffee farms and pastures, and in suburban yards. It occurs from sea level to high montane regions. The highest ant record I have for Costa Rica, a collection at 3000 m near Villa Mills, is Labidus coecus. The species is almost entirely subterranean, sometimes at considerable depth... In the study of army ants, most of the attention has focused on the large epigaeus species in the genus Eciton. But the highest density and most ecologically important army ants may turn out to be L. coecus." Crawley (1916) noted this underappreciation, writing that L. coecus in Guyana is "a common species, but owing to its habit of burrowing beneath the surface of the soil it is not frequently observed." Smith (1965) wrote that L. coecus: "is a native and widely distributed species, ranging from Oklahoma and Arkansas to Texas and Louisiana, and south to Argentina." Recently, however, Wittenborn and Jeschke (2011) included L. coecus on their list of exotic ant species established in North America. Here, we examine the biogeography of L. coecus. We were particularly interested in evaluating evidence concerning whether or not L. coecus has established any exotic populations in North America or elsewhere.

Material and methods
Using published and unpublished records, we documented the known range of L. coecus. We obtained unpublished site records from museum specimens in the collections of Louisiana State Arthropod Museum (LSAM); the Museum of Comparative Zoology (MCZ), and the Smithsonian Institution (SI). We obtained unpublished site records of L. coecus from C. Sanabria (Colombia), A.J. Pérez-Sánchez (Margarita), and E. Mendoza (El Salvador). In addition, we used on-line databases with collection information on specimens by the Field Museum, Antweb (www.antweb.org), and the Global Biodiversity Information Facility (www.gbif.org). JKW collected L. coecus in Tobago and El Salvador.
We obtained geo-coordinates for collection sites from published references, specimen labels, maps, or geography web sites (e.g., earth.google.com, www.tageo.com, and www.fallingrain.com). Published records usually included collection dates. In a number of cases, publications did not include the collection dates for specimens, but we were able to determine the approximate date based on information on the collector's travel dates. For example, Forel (1899a) reported L. coecus from Colombia collected by Princess Theresa of Bavaria, who took part in a scientific expedition to South America in 1898.

Results
We compiled and mapped >650 site records (Fig. 1), documenting the earliest known record for 27 geographic areas (countries, US states, and major West Indian islands), including three for which there are no previously published records: Margarita, Tobago, and Trinidad (Table 1).
In the US, the vast majority of L. coecus records came from Texas. O' Keefe et al. (2000) listed records of L. coecus from 67 counties in Texas. We found records from six additional counties in Texas, primarily from caves: Burnet, Coryell, Medina, Menard, Montague, and Val Verde (Reddell and Cokendolpher 2001, Calixto 2008, Cokendolpher et al. 2009. Watkins (1976) included a distribution map for L. coecus showing continuous range from Buenos Aires, Argentina to Oklahoma, Arkansas, and Louisiana with gaps around Uruguay and (inexplicably) Nicaragua.
We found no published site records for L. coecus in Oklahoma. Smith (1935) wrote that in Oklahoma, L. coecus "may range into the extreme southern part of the state." Later, Smith (1947) listed L. coecus as occurring in Louisiana, Oklahoma, and Texas, but gave no site information. Smith (1965Smith ( , 1979 then listed L. coecus from Arkansas, Louisiana, Oklahoma, and Texas. We found only one L. coecus specimen from Oklahoma, collected at a black light in Delaware County (1971; C. M. Smith; LSAM).
We found no specific site records for L. coecus in Uruguay. Fernández and Sendoya (2004) listed L. coecus in Uruguay citing Borgmeier (1955), Watkins (1976), and Palacio (1999) as their source of information on this species, but none of these references recorded L. coecus in Uruguay. Zolessi et al. (1989) did not include L. coecus in their summary of the ants of Uruguay. Nonetheless, we mapped the Uruguay record to Artigas in northern Uruguay, near records in southern Brazil. It seems almost certain that L. coecus does occur in Uruguay, given its presence in surrounding parts of Brazil and Argentina.

Problematic records
Antbase coded one L. coecus record as from Espiritu Santo, Vanuatu, but the specimen was actually from Espiritu Santo, Brazil. Antbase previously listed a record of L. coecus from New Mexico, but this turned out to be a misidentification (C. Moreau, pers. comm.).

Discussion
Labidus coecus has now been reported from 27 geographic areas, including 20 countries in Central and South America (all except Chile), three West Indian islands (Margarita, Tobago, and Trinidad), and four US states (Arkansas, Louisiana, Oklahoma, and Texas). The range of L. coecus appears to be essentially continuous, from Buenos Aires, Argentina (~34.6°S ;Borgmeier 1955) in the south to Delaware County, Oklahoma (~36.6°N; see Results) in the north. The three West Indian islands with L. coecus populations are all continental shelf islands that were connected to South America during   (53), Mexico (4), Venezuela (1), and Peru (1) (Kempf 1961, Reddell and Cokendolpher 2001, Cokendolpher et al. 2009). The more stable microclimate of caves may allow L. coecus to extend its range into regions with otherwise inhospitable climates. Given its extremely broad ecological tolerance, it is unclear why L. coecus populations in the southeast US are not known east of Louisiana, when there would appear to be suitable habitat for this species below 31°N all along the Gulf coast of Alabama and Mississippi and into Florida. Wittenborn and Jeschke (2011) included L. coecus in a list of 93 exotic ant species established in North America. The populations of L. coecus, however, appear to be continuously distributed from Argentina to the southern US. We know of no evidence that even suggests that L. coecus has any exotic populations. In addition to L. coecus, Wittenborn and Jeschke (2011) appear to have misclassified as exotics numerous other ant species that are actually native to North America. For example, Gnamptogenys hartmani (Wheeler), Leptogenys elongata (Buckley), and Pachycondyla harpax (Fabricius) all have distributions in the southern US that appear to be the northern end of continuous native ranges and give no indication that these species are exotic to North America. Other species that Wittenborn and Jeschke (2011) most likely misclassified as exotics include Cephalotes varians (Smith), a widespread arboreal species known from Cuba, the Bahamas, and Florida (de Andrade and Baroni Urbani 1999) and Leptogenys manni (Wheeler), a species endemic to Florida (Trager and Johnson 1988). For more than 20 additional ant species, Wittenborn and Jeschke's (2011) classification as exotic in North America is questionable. It seems hazardous research protocol to list species as exotic without documenting the source of this classification. In addition, there is some danger that if native species are erroneously considered to be exotics, they may be treated as such and exterminated, rather than valued and protected.

studies of laboulbeniales (Fungi, Ascomycota) on Introduction
The order Laboulbeniales (Ascomycota: Laboulbeniomycetes) consists of microscopic ectoparasites of Arthropoda, mostly true insects. Within the subphylum Hexapoda representatives of nine orders are known as hosts (Weir and Hammond 1997): Blattodea, Coleoptera, Dermaptera, Diptera, Hemiptera, Hymenoptera, Mallophaga, Orthoptera, and Thysanoptera. [Note that the termites, previously ranked in the order Isoptera, were recently included in the order Blattodea, based on phylogenetic data (Beccaloni and Eggleton 2013).] In addition, Acari (subphylum Cheliceriformes) and Diplopoda (subphylum Myriapoda) are known to host Laboulbeniales (Haelewaters et al. 2012, Weir andHammond 1997). Laboulbeniales are unusual in their determinate growth pattern and lack of hyphae. Diversity in the group is largely underexplored and many questions related to the taxonomy and biology of these fungi remain unresolved.

Methods
The study site spans somewhat less than 1 km 2 , situated east of Maastricht (Limburg, the Netherlands) near the border with Belgium. This area has a rolling landscape and calcareous soil. Many different habitats were sampled: calcareous grassland, thicket, moist forest, forest edge, felling area, agricultural field edge, and hollow road.
Pitfall traps were filled with a formol solution (3%) or with a saturated salt solution. A lid was placed above each trap to exclude rainfall. Pitfall trapping was performed during three inventories for which in total 424 pitfall traps were placed in 29 series: (i) a year-round inventory of the insect fauna with nine trap series (each with five pitfall traps) at a railroad verge with semi-natural grassland and thicket, (ii) a twomonth-long inventory with eleven trap series (each with five pitfall traps) of the arthropod fauna of a nature restoration parcel with a species-rich hay-meadow (dominated by Arrhenatheretum elatioris Braun-Blanq.), and (iii) a four-day-long inventory of nine different habitats (nine traps series, one series per habitat, each with 36 pitfall traps) during a student course. For an overview of the study area, see Figure 1.
Screening for thalli of Laboulbeniales was done at 45× magnification with a Euromex Z-1740H stereomicroscope (Arnhem, the Netherlands). Infected ants were sent to Harvard University for study of their associated fungi. Thalli were carefully removed from the host integument using a Minuten Pin (BioQuip #1208SA) and embedded in Amann solution (Benjamin 1971) or PVA Mounting Medium (BioQuip #6371A). Cover slips were ringed with transparent nail varnish. Morphological analyses, measurements, and identifications were done using an Olympus BX40 light microscope with Olympus XC50 digital camera and MicroSuite Special Edition software 3.1 (Soft Imaging Solutions GmbH). Voucher specimens are deposited at the Farlow Herbarium, Harvard University (FH).
Chi-square tests were performed to analyze whether infection rate of the infected ant species was different from a hypothesized even distribution of Rickia wasmannii presence over the infected species. In social insects, it sometimes seems better to analyze the number of infected nests instead of the number of infected workers. However, the infected Myrmica species are very abundant in our study area, are polygynous and occur in clusters of mini-populations. They are hardly territorial and contact between workers of different nests is likely (Garnas et al. 2007, Wilson 1971. Each pitfall trap (and trap series) thus likely samples ants from many nests. Our sampling effort with 424 pitfall traps is so widespread over the study area and combines workers from so many nests, that we performed our test on worker individuals.

Results
Twenty-seven ant species were recorded during this study (Table 1). Only three species in the genus Myrmica bore thalli of Rickia wasmannii.
Within the current study, the highest parasite prevalence was found on Myrmica sabuleti: 38% (n = 3,876; Figure 2). For the other two species, parasite prevalence was considerably lower: 11% on M. scabrinodis (n = 643) and 0.55% on M. ruginodis (n = 182). These infection rates differ significantly when an even proportional distribution of Rickia over the three species is presumed, with M. sabuleti a significantly higher infection rate as would be expected by chance (observed number of infected ants: 1479; expected: 1279; χ 2 =46.82, df=2, p<0,001), and a significantly lower infection rate for both M. scabrinodis (observed number of infected ants: 71; expected: 212; χ 2 =140.11, df=2, p<0,001) and M. ruginodis (observed number of infected ants: 1; expected: 60; χ 2 =68.64, df=2, p<0,001). When we look at Rickia prevalence over the 29 traps series instead of at the level of individual workers, the same pattern emerges: infected M. sabuleti workers in 15 series, infected M. scabrinodis in five series, and infected M. ruginodis in only one series.
Only workers were found infected; neither gynes nor males were found infected, not even in the highly infected Myrmica sabuleti. Myrmica ruginodis and M. scabrinodis are reported as hosts of Rickia wasmannii in the Netherlands for the first time.

Discussion
Rickia wasmannii is mentioned in the literature in the literature to occur on nine host species (Table 2), most of which are widespread across Europe (Radchenko and Elmes 2010). It is reported from sixteen European countries, and expected to occur in Belgium, Denmark, Ireland, and Portugal (Espadaler and Santamaría 2012, Haelewaters 2012, Witek et al. 2014).

Ant species with Rickia infection
Santamaría and Espadaler (2014) mention the "high host phylogenetic specificity" of Laboulbeniales, but caution is needed when interpreting support for this assertion. Although specific to the genus Myrmica, host species of Rickia wasmannii belong to two clades or so-called species groups that are not phylogenetically closely related: rubragroup and scabrinodis-group (Jansen et al. 2010).
The Palearctic species in the genus Myrmica are classified into 17 taxonomic species groups based on their morphology (Radchenko and Elmes 2010), three of which are represented in our study: the rubra-group, encompassing M. rubra and M. ruginodis; the scabrinodis-group, with M. rugulosa, M. sabuleti, and M. scabrinodis; and the schencki-group, to which (in most parts of Europe) only M. schencki belongs. The monophyly of these morphological species groups was confirmed using molecular data (Jansen et al. 2010). So far, only species of the rubra-group and scabrinodis-group have been found with Rickia wasmannii infection. Our research confirms this finding: although six Myrmica species were screened (n = 6,314), Rickia wasmannii was only present on M. ruginodis, M. sabuleti, and M. scabrinodis (Table 1). Several researchers screened multiple species of Myrmica in Hungary/Romania (Tartally et al. 2007), Slovakia (Bezdĕčka and Bezdĕčkova 2011), and the Czech Republic (Bezdĕčkova and Bezdĕčka 2011), but found only species in the scabrinodis-group infected by R. wasmannii. In addition, our study in the Netherlands revealed a single lightly infested worker in the rubra-group (M. ruginodis). In all four studies, R. wasmannii was found only in the rubra-and scabrinodis-groups.
Inadequate sampling of potential hosts may (partly) explain this pattern, although recently in Europe several studies have been conducted on Laboulbeniales on Myrmica, including various studies in which different species groups were sampled and screened. To avoid taxon-sampling errors in these kinds of observation, we suggest that systematic collections in natural history museums be screened for Rickia wasmannii on Myrmica. Screening museum collections previously has yielded important contributions on patterns of host utilization (Weir and Hammond 1997: 80,000 insects screened) and the distribution of Hesperomyces virescens Thaxt. (Haelewaters et al. 2014: 4,000 ladybirds screened).
Host shift could be another explanation for the restricted presence of R. wasmannii on two species groups that are not sister clades. Host shifts have been suggested for morphologically similar Laboulbenia species between Cicindelinae and other Car- abidae living in the same habitat (Arndt et al. 2003, Rossi 2011), and De Kesel and Haelewaters (2014 provide morphological and ecological data to support the hypothesis that a species of Laboulbenia shifted between Carabidae and Staphylinidae. To explain the extremely small size of Rickia lenoirii on Messor spp. ants (Hymenoptera, Formicidae), Santamaría and Espadaler (2014) suggest that Laboulbeniales can shift between myrmecophilous Acari and their ant hosts.

Geographical variation
Parasite prevalence (= infection frequency) is often used to quantify differences in populations of Laboulbeniales in a given host community (De Kesel 2011. It is beyond doubt that in our study area Myrmica sabuleti is the main host of Rickia wasmannii, considering the actual prevalence of R. wasmannii on M. sabuleti workers infected, compared to the low parasite prevalence on M. ruginodis and M. scabrinodis (Table 1).
In our study site, parasite prevalence was high in the scabrinodis-group (Table 1). Myrmica sabuleti and M. scabrinodis occur in other habitats; the former species gener-  (Bezdĕčka and Bezdĕčkova 2011;Bezdĕčkova and Bezdĕčka 2011;Csata et al. 2013;Espadaler and Santamaría 2012;Haelewaters 2012;Tartally et al. 2007;Witek et al. 2014), completed with the current findings from the Netherlands. *In Hungary and Romania, M. scabrinodis is the most common host of R. wasmannii (Csata et al. 2014;Tartally et al. 2007 ally lives in drier areas than the latter. While in our study area in the province Limburg both species live in sympatry, we observed noticeable differences in occurrence and parasite prevalence. In the rubra-group, only one specimen of M. ruginodis was found with Rickia wasmannii. The fact that we did not find any infected M. rubra out of 974 workers raises questions, since this species is the second-most commonly found infected species in Europe after M. scabrinodis, with examples in Austria, Germany, Luxembourg, Romania, and Switzerland (Espadaler and Santamaría 2012, Table 2). Also in Romania, so far, infection by Rickia wasmannii only occurs in the rubra-group and scabrinodis-group (Csata et al. 2013, Tartally et al. 2007). In Romania, Myrmica scabrinodis often is the only or main host; in a single population R. wasmannii was found only on M. rubra, although thalli were also observed in populations of M. scabrinodis and M. slovaca; M. gallienii and M. rubra; and M. ruginodis (Csata et al. 2013).
Myrmica scabrinodis is the main host in Romania (studied areas, Csata et al. 2013), as is M. sabuleti in our studied site in the Netherlands. These observations suggest (1) that R. wasmannii has a considerable host choice plasticity within the genus Myrmica and (2) that infection rates of Rickia wasmannii show geographical variation, potentially with different dominant host species across regions. It is not known what factors cause this geographical variation. Future research in the Netherlands is needed to confirm that M. sabuleti is the main host in a wider region.

Introduction
The solitary, ground-dwelling wasp Cerceris fumipennis Say currently is being employed as a tool for biosurveillance of pest Buprestidae in eastern North America as well as for the documentation of regional buprestid diversity (reviewed by Swink et al. 2013, Careless et al. 2014. Any observations that contribute to more efficiently using this unique system is therefore of interest and value. Here we focus on prey carriage and paralysis by the wasp. Current literature offers mixed reports of prey carriage by Cerceris fumipennis. Although there is general agreement that the beetle is carried in the wasps' mandibles with its head forward and venter facing up, where the beetle is grasped, and the involvement of the wasp legs in supporting the prey varies. Mueller et al. (1992) indicate that C. fumipennis appears to use just legs to hold larger buprestids while in flight, but Careless (2009) reports that an antenna of large beetle prey is grasped in the mandibles, with the wasp fore-and mid-legs embracing the body. Available images of C. fumipennis with prey are of little help, because they typically show the wasp perched on vegetation or the ground, or on final approach with her landing gear down. There are a few exceptions. Careless et al. (2009: fig. 23) picture a female in flight carrying a prey item about half her length. The wasp metasoma is extended, the thorax of the prey beetle is under her head, and the fore and middle legs are supporting the prey with the hind legs flexed and uninvolved. In video footage of a wasp hovering near her nest entrance with a large prey item (Walton 2011), it is fairly clear that all three sets of legs embrace the prey, but involvement of the mandibles is not visible. Mueller et al. (1992) reported that smaller beetle species, including those in the genus Agrilus, are clasped about the mid-thorax with the wasp mandibles. In the course of six seasons of working with Cerceris fumipennis, however, the authors (CAN and WGS) independently noted that females coming back to the nest carrying small, Agrilus-sized beetles often can be recognized in flight by the curvature of the wasp's metasoma (Fig. 1d). Investigation into the basis of this flight posture led us to Krombein (1981), who described a "buprestid clamp" on the fifth metasomal sternite of Cerceris females in the bupresticida group, including C. fumipennis. He described it as a median concavity with an erect lamella at the posterior margin (Fig. 1a-c). He goes on to state that the female "grasps the head of the buprestid in her mandibles, curves the tip of her abdomen beneath and forward so that the abdominal apex of the buprestid rests in the concavity and is prevented from slipping by the erect flange. This clamp functions only with long slender buprestids such as Agrilus species." He found it a pleasing example of the correlation between structure and function.
Krombein supplied the foundation for the unique flight posture we observed, but it seemed unusual that a female transporting a small paralyzed beetle in her mandibles had need to stabilize such a light load by means of a morphological structure specific to the purpose. Here we suggest that the basis of specialized carriage of small prey by Cerceris fumipennis is that small beetles, including most Agrilus species, often are not successfully paralyzed at their collection site.

Methods
During biosurveillance with Cerceris fumipennis in North Carolina during 2014, a subset of wasps captured with prey from seven widely scattered nesting aggregations were analyzed. A total of 258 females carrying prey were captured in a sweep net and the activity level of the prey item rated as paralyzed (incapable of locomotion) or not paralyzed (capable of escape). The beetle prey were then transported on ice to the laboratory where body length was measured using digital calipers (Mitutoyo Absolute Digimatic caliper -0.01mm). Individual beetles were measured 3×, and the average recorded. Beetles were identified by W.G. Swink and J.P. Basham (Tennessee State University).
Overall, 27 Agrilus (nine species) were captured; in three species, both paralyzed and active individuals were collected from foraging wasps (Table 1). Just one individual of A. pensus was collected overall, and it was active when collected. The 27 Agrilus captured ranged from 4.9 mm in length (A. subrobustus) to 10.7 mm (A. ferrisi). There was no significant difference in the length of Agrilus that were paralyzed vs. those that were not (t = 0.31, P = 0.76; two-sample t-test). It is notable that the smallest Agrilus captured (A. subrobustus Saunders) was paralyzed. The single active Brachys ovatus measured 5.4 mm. Krombein's (1981) proposal that the buprestid clamp on Cerceris fumipennis is specialized for the transport of Agrilus, combined with the inconsistent paralysis of Agrilus spp. reported here suggests that the two are related. Stinging behavior as well as "the state of the stung" is known to vary considerably with context (Steiner 1986), and it is likely difficult for C. fumipennis to manipulate small buprestids into a suitable position for stinging at the capture site. All Buprestidae are well sclerotized, and the vulnerable coxal membranes ) of small beetles offer a miniscule target for insertion of the stinger. All buprestids furthermore exhibit death feigning (thanatosis) as a defense mechanism; they typically respond to disturbance by retracting their appendages and becoming quiescent. This allows the wasp to carry a compact and motionless beetle into its nest, even if stinging was unsuccessful (Careless 2009). We suggest that the buprestid clamp in C. fumipennis functions primarily to prevent the elytra from opening in small prey that may or may not be successfully paralyzed. This applies to those in the genus Agrilus (as suggested by Krombein 1981), as well as to other small species with an abdominal tip narrow enough to fit in the concavity of the clamp, such as Brachys ovatus and perhaps the alternative prey Neochlamismus bebbianae Brown (Chrysomelidae) (references in Careless et al. 2014). The latter is a small, heavily sclerotized beetle that has been taken nine times from C. fumipennis in North Carolina, and none to date have been paralyzed (WGS, pers. obs.).

Discussion
Medium to large beetles also may be less than completely paralyzed prior to transport, but the leg embrace by the wasp assures their immobility during flight. Once in the nest, the wasp is free to assure paralysis of the prey at her leisure. Prey temporarily stored in the main burrow, prior to placement in the brood cells, continue to vary in their degree of paralysis; an Agrilus bilineatus was observed crawling out of the nest by Hook and Evans (1991). However, beetles are stung (or re-stung) prior to final placement in the brood chamber Hook and Evans 1991), and prey collected from Cerceris fumipennis brood cells are always "rather thoroughly paralyzed" (Kurczewski 1984, Hook andEvans 1991).
Prey carriage plasticity in Cerceris fumipennis is related to the remarkable range of both size and shape of their buprestid prey. Careless (2009) reported that the beetles he collected were 4.9-22.3 mm in length, and those taken from C. fumipennis in North Carolina in 2014 exhibit a similar range (4.9-21.3 mm; n=258; Nalepa and Swink, unpublished data); shape varies from the elongate, parallel-sided Agrilus to the broadly oval species of Buprestis. In medium to large beetle prey, an anterior body part of the beetle is clasped by the wasp mandibles and the number of legs involved in load support likely depends on prey length, their weight in relation to the center of gravity during wasp flight, and perhaps the degree of beetle paralysis. The load needs to be not only secure, but balanced during transport. Small prey are grasped with the mandibles, and secured via the buprestid clamp. As a whole, then, C. fumipennis falls into two established categories of prey carriage mechanisms depending on prey size: Mandibular 3, where the prey is held in mandibles and supported in flight with the legs, and Abdominal 2, with prey carried in flight supported by structural modifications of the apical metasomal segment (Evans 1962, O'Neill 2001.
Species composition of the buprestid prey of Cerceris fumipennis varies with geographic location and the plant species composition surrounding nesting aggregations.
For example, of the 310 beetles taken from C. fumipennis nests by Evans (1971) in New York State, 84.5% were Agrilus spp., while just 8.2% of the 466 beetles collected from C. fumipennis in the Goldsboro area of North Carolina over four years were in that genus (Swink et al. 2014). Nonetheless, C. fumipennis has specific morphological and behavioral adaptations to secure and transport Agrilus, suggesting that this buprestid genus is an evolutionarily significant prey item of the wasp. Such specific adaptations to prey transport are exhibited by additional Cerceris species that hunt Buprestidae (Krombein 1981), as well as by the weevil hunting C. halone (Byers 1978).

Conclusions
The genus Agrilus contains mostly small, elongate species (4-13 mm -Paiero et al. 2012), many of which are economically important and thus the primary target of biosurveillance programs. Collections from Cerceris fumipennis can be biased against Agrilus, however, not only because of the speed of females laden with small prey, but also because of their ability to pass though, unimpeded, the plastic "Careless collars" designed to slow or stop females carrying larger beetles (see Careless et al. 2014: Fig. 1b). The latter drawback can be overcome, at least in part, by placing the hole in the collar slightly askew over the burrow entrance (Careless et al. 2014). Here we suggest that the bias can be further mitigated by expanding the search image for prey-laden incoming wasps. Cerceris fumipennis females carrying Agrilus and other small buprestid species can be recognized by their characteristic curved posture while in flight, even if the prey item is not visible. Attempts to capture these females in a sweep net prior to their arrival at the nest can help fine tune the biosurveillance system and increase the odds of detecting pest Buprestidae.

Colemanus keeleyorum (Braconidae, Ichneutinae s. l.): Introduction
Herein we describe Colemanus keeleyorum gen. n. et sp. n. (Braconidae: Ichneutinae s. l.) from the Eocene Green River Formation in the western United States. The Green River Formation represents one of the best documented ancient lake systems and has offered a particularly well preserved window into Eocene fauna. The formation is best known for fossil fish (Conrad et al. 2007), but other key discoveries include the oldest known bats: Icaronycteris index Jepsen, 1966 andOnychonycteris finneyi Simmons et al., 2008. Recent collecting efforts by David Kohls (Colorado Mountain College) and colleagues have accumulated more than 120,000 specimens, including more than 2,000 insects. One specimen, a wasp collected in 2005 in the Parachute Member of the Piceance Creek Basin (northwestern Colorado), caught the attention of Jim Whitfield (University of Illinois) due to its resemblance to members of a large braconid assemblage called the microgastroid complex.
The microgastroid complex is a well-supported rapid radiation (e.g., Whitfield and Lockhart 2007, Whitfield and Kjer 2008, Murphy et al. 2008, Sharanowski et al. 2011 well-known for its association with host-immunosuppressing polydnaviruses (e.g., Whitfield 2000). Overall, the relationships of subfamilies within the complex are poorly understood due to the rapid nature of their diversification (Whitfield andLockhart 2007, Whitfield andKjer 2008), but two relationships are clear. First, Cheloninae is sister to all other subfamilies (Whitfield 1997, Dowton and Austin 1998, Dowton et al. 1998, Belshaw et al. 2000, Banks and Whitfield 2006, Murphy et al. 2008, Sharanowski et al. 2011. Second, Ichneutinae s. l., usually excluded from the microgastroid complex because members are not known to contain polydnaviruses, is sister to the microgastroid complex (Quicke and van Achterberg 1990, Belshaw et al. 2000, Belshaw and Quicke 2002, Dowton et al. 2002, Murphy et al. 2008, Sharanowski et al. 2011. Ichneutinae has received considerable attention and endured a chaotic taxonomic history, despite its relatively small size of approximately 100 species (Sharkey 1994, Yu et al. 2005. Three lineages are recognized and traditionally assigned to the tribal level (Proteropini, Ichneutini, and Muesebeckiini); however, tribal designation has been disputed Sharkey 1994Sharkey , 1996. Although normally treated as a single subfamily, phylogenetic analysis often renders Ichneutinae paraphyletic (e.g., Quicke and van Achterberg 1990, Belshaw et al. 2000, Dowton et al. 2002, Sharanowski et al. 2011. The most rigorous of these analyses, which investigated multiple ribosomal and nuclear genes, recovered Proteropini as sister to microgastroids and Ichneutini as sister to that complex (Sharanowski et al. 2011). The affinities of the highly-derived muesebeckiines remain unknown as they were excluded from that analysis.
Placing the fossil within the Ichneutinae s. l. + microgastroid complex is difficult due to its resemblance to several lineages. Specifically, the fossil resembles Cardiochilinae because the last abscissa of the fore wing radial sector (3RSb) is recurved, although unlike many cardiochilines this vein remains tubular as it reaches the wing margin; and it resembles Proteropini (Ichneutinae s. l.) because the first abscissa of the fore wing media (1M) is evenly curved. Some Cheloninae also share these characters, but are readily differentiated by the presence of a metasomal carapace.
To determine the fossil's affinity, we placed it on a phylogenetic hypothesis for Braconidae (Sharanowski et al. 2011) using a parsimony criterion (tree length) of morphological characters. The dataset was expanded by adding taxa superficially resembling the fossil (Bohayella and Schoenlandella (Cardiochilinae); Helconichia (Proteropini)). The method implemented herein has the benefit over a standard analysis of easily comparing alternative placements of the fossil.

Materials and methods
The photographs of Colemanus gen. n. (Fig. 1) were taken with a JVC KY-F75 3CCD digital camera and prepared with Archimed™ by Microvision Instruments. Reconstructions (Fig. 2) were created by tracing these photographs in Adobe Illustrator CS4, and by editing in Adobe Photoshop CS4. Unfortunately, the metasoma and legs did not preserve during fossilization (or are obscured by the rock matrix), and the head is crushed, offering no codable characters for these regions. The wings are remarkably preserved, complete with wing setae, whereas the dorsal mesosoma and antennae are moderately well preserved.

Characters
States Code Source they were poorly preserved. These characters are as follows: the degree of notauli sculpturing posteriorly, the medial carina on the lateral scutellum, and the carinae on the lateral metanotum. The remarkably preserved wings provided most detail and were the principle units used in morphology. Both wings showed some bending, so both wings were used to create a composite hypothesis for wing veins (Fig. 2B). Areas of the wing where unbiased reconstruction were impossible include the apparently non-tubular region surrounding (RS+M) b (Fig. 1C), the apical non-tubular limits of the fore wing media and cubitus, and the apical non-tubular limits of the hind wing radial sector and media. Noteworthy is that (RS+M) b appears to be vertical, which is a rare character for all braconids, although this region is difficult to interpret. Also, the last abscissa of the radial sector (3RSb) is recurved, lending the resemblance to Cardiochilinae and some Cheloninae, although 3RSb in cardiochilines is not tubular as it reaches the wing margin and chelonines have a metasomal carapace.
Despite the great condition of the antennae, no characters could be extracted except flagellomere number (33-34), which itself is inconclusive due to the indiscernible basal antennomeres. Regardless, there are more antennomeres than in other described proteropines (24-31). However, given the variability of this character across Ichneutinae s. l. (12-38), this difference is not considered informative at the level of our analyses and is not included in the matrix.
Morphological phylogenetics: Tree lengths representing placement of Colemanus at each possible node can be viewed in Figure 3, across the three possible relationships of Proteropini (A-C). Without the addition of Colemanus, total tree lengths varied (33-35) depending on the arrangement of Proteropini (boxed values in Fig. 3). Adding Colemanus raises total tree lengths to a minimum of 36 (when placed sister to Helconichia+Muesonia, given the topology in Fig. 3A) and a maximum of 44 (when placed sister to Cardiochiles, given the topology in Fig. 3C). For each topology, the most parsimonious placement of Colemanus is within Proteropini.
We conclude from these findings that Colemanus should be placed within Proteropini, although given the uncertainty of proteropine relationships, exact placement within this group is not yet feasible. The fossil contains character states that do not fit within any current genus. Therefore, we suggest placement within a new genus, Colemanus.  Di agnosis. There are several similarities between Colemanus and other braconids, specifically Cardiochilinae and Cheloninae. Like Colemanus, some Cheloninae have a recurved 3RSb and an evenly curved 1M. However, chelonines possess a metasomal carapace. Colemanus contortus comb. n. lacks a carapace (Brues 1933) and although the metasoma of C. keeleyorum sp. n. is not completely visible, it seems apparent that it also lacks a metasomal carapace (Fig. 1A). Thus, the lack of a metasomal carapace distinguishes Colemanus from Cheloninae. Colemanus also resembles many Cardiochilinae in having a recuved 3RSb and heavily sculptured mesosoma, but can be readily distinguished by 3RSb remaining tubular as it reaches the wing margin.
Colemanus can be distinguished from other Ichneutinae s. l. by the presence of a curved 3RSb; fore wing 1cu-a curved downward, not angled toward wing margin; hind wing M+Cu positioned in the posterior half of the wing; and a heavily sculptured mesosoma.
Etymology. Named for bodybuilder Ronnie Coleman, who was famous for his back; referring to the robust and sculptured nature of the mesosomal dorsum.
Description. Holotype (n = 1): body length 9 mm (estimated due to incomplete metasoma); sex unknown. Head (Fig. 1A) 1.4 mm long and 1.7 mm wide. Antenna 7.2 mm long with 33-34 flagellomeres. Mesosoma (Figs 1B,2A) 3.8 mm long and 3.3 mm wide; robust and heavily sculptured; notauli deeply crenulate; scutellum with crenulate depression medio-posteriorly; side of scutellum deeply hollowed with crenulate carinae and bordered by prominent carinae; metanotum with median, raised, rectangular tubercle having radiating carinae; scutellar sulcus with carinae; propodeum with areolate sculpturing. Wings (Figs 1A,C,2B) with last abscissa of fore wing radial sector (3RSb) recurved and tubular as it reaches the wing margin; (RS+M)b short, nearly vertical; 1cu-a originating far distal to M and curving downward, thus not angling toward wing apex; fore wing M curved; tubular portion of 1a long; C+SC and R closely fused; second abscissa of RS strongly curved; parastigma well-developed; hind wing M+Cu longer than first abscissa of M; hind wing r-crossvein absent; hind wing M+Cu in posterior half of wing; 1A strongly developed.
Remarks. Wings, antennae, and dorsal mesosoma are overall well-preserved; metasoma and legs either did not completely fossilize or are obscured by the rock matrix; head is crushed. Etymology. Named for Dr. Jack and Flo Keeley, who, together with their daughter and her husband (first author's mother & father), were largely responsible for the first author's pursuit of the natural sciences. Colemanus contortus (Brues, 1933), comb. n.
The material Brues (1933) examined is part of the Königsberg collection. Most of this collection was rescued and is housed at the Geowissenschaftliches Museum (Göttingen, Germany), though some of Brues's collection, which included some of the Königsberg collection, remains at Harvard University, his old institution. However, neither the Museum of Comparative Anatomy (Harvard), nor the Geowissenschaftliches Museum could locate Brues's 1933 material, including Ichneutes contortus. The problem is exacerbated by the fact that Brues did not include catalog numbers in his description or any other information regarding deposition. Therefore, the holotype of I. contortus should be considered lost.
Key to Colemanus gen. n.

Introduction
The genus Labania Hedqvist is a small doryctine group restricted to the Neotropical region and is currently composed of four species. This genus was erected by Hedqvist (1963) based on a single species from Honduras, L. straminea Hedqvist. In the description, Labania was placed within the morphologically heterogeneous subfamily Hormiinae, but it was subsequently transferred to the Doryctinae in a tribal reclassi-fication of this subfamily (Belokobylskij 1992). To date, only three additional species of Labania have been described from Costa Rica (Marsh 2002), L. hansoni Marsh, L. minuta Marsh and L. prolata Marsh, with the first two being reared from aerial root galls on Ficus citrifolia Mill. and leaf galls on F. colubrinae Strandl., respectively. In his tribal reclassification of the Doryctinae, Belokobylskij (1992) placed Labania as the single member of the tribe Labaniini based on several putative apomorphic features. More recently, however, three molecular phylogenetic studies consistently recovered a clade exclusively composed of genera whose species are associated with galls from different plant families, including Labania (Zaldivar-Riveron et al. 2007, 2014. In the most recent of these studies (Zaldívar- , the authors described three new gall-associated doryctine genera whose species with rearing records were all associated with Ficus species. One of them, Ficobolus Belokobylskij & Zaldívar-Riverón, was recovered as sister to Labania; the latter genus was represented in that study by an undescribed species from Costa Rica. In this work, this new species of Labania, which was reared from aerial roots on F. obtusifolia Kunth is described, and observations that suggest that this new species is phytophagous are provided. An updated key (adapted from Marsh 2002) to the five described species of Labania is given.

Material and methods
All specimens of the new species of Labania described here were reared from a single tree of F. obtusifolia (see details below). Specimens belonging to two of the remaining species of the genus were also examined, as well as digital photographs of the holotype of L. straminea, deposited at National Museum of Natural History (Washington, D.C., USA). The type material of the new species is deposited in the following collections: Colección Nacional de Insectos, Instituto de Biología, Universidad Nacional Autónoma de México (CNIN IB-UNAM); Museo de Zoología, University of Costa Rica, San José, Costa Rica (MZCR); Zoological Institute of the Russian Academy of Sciences, St. Petersburg, Russia (ZISP); and National Museum of Natural History, Washington, DC (NMNH).
The terminology employed in this work for the morphological features and measurements follows Belokobylskij and Maetô (2009). The wing venation nomenclature follows Belokobylskij and Maetô (2009) and  in parentheses. Photographs were taken with a Leica IC 3D digital camera mounted on a Leica® MZ16 microscope and using the Leica Application Suite® imaging system.

Taxonomy
Mesosoma: length 1.6-1.7 times its height. Median lobe of mesoscutum distinctly and highly convex (lateral view), slightly protruding forward and rounded long as first tergite of metasoma, 0.40-0.45 times as long as mesosoma, 0.18-0.22 times as long as fore wing.
Sculpture and pubescence: head entirely smooth, sometimes frons near antennal sockets with several curved striae. Mesoscutum smooth, with rugulosity in short and small medioposterior area. Scutellum mainly smooth, finely striate laterally. Mesopleuron mostly smooth. Propodeum widely rugose-striate, with all areas distinctly delineated by carinae; basolateral areas short, semi-rounded, mainly smooth; areola wide, about as long as maximum width; basal carina 0.2-0.3 times as long as basal fork of areola. Hind coxa and femur smooth. First tergite mainly striate with rugosities, finely reticulate to almost smooth between dorsal carinae. Remaining tergites entirely smooth. Vertex with numerous, dense and semi-erect pale setae. Mesoscutum mainly glabrous, with dense and erect pale setae arranged in almost single lines along notauli and laterally. Mesopleuron medially rather widely glabrous. Hind tibia dorsally with dense, relatively long and semi-erect pale setae, its length 0.5-0.8 times maximum width of hind tibia.
Colour: head entirely light reddish brown or sometimes yellowish brownish, ocellar triangle often black. Antenna black, dark reddish brown basally, two basal segments yellowish brown. Palpi pale brown. Mesosoma almost reddish black, with light brown on narrow stripe along lower margin of pronotal sides and along area of precoxal sulcus. Metasoma mainly dark reddish brown, dorsal basal and apical parts black. Fore and mid-  Marsh -Transverse diameter of eye 2.0-2.2 times wider than temple (Fig. 1C). Eye covered with short and sparse setae. Malar space 0.35-0.40 times height of eye (Fig. 1D). Propodeum mainly rugose-striate, smooth only in basolateral areas (Fig. 1C). Discoidal (first discal) cell sessile (Fig. 1F). Dorsal carinae of first metasomal tergite almost complete (Fig. 1E)