Revision of Therophilus s.s. (Hymenoptera, Braconidae, Agathidinae) from Thailand

Based on a phylogenetic analysis, the limits of Therophilus (Hymenoptera: Braconidae: Agathidinae) are redefined and restricted to a small proportion of the previously included species. Those species belonging to the world fauna are listed and the species from Thailand are revised. Forty-four species are assigned to the genus including 11 new species, i.e. T. anuchati, T. apichati, T. areeluckae, T. boonthami, T. chiangmaiensis, T. kwanuiae, T. songrami, T. sukpengae, T. wannai, T. wongchaii, T. wongwani. A dichotomous key to species is presented; links to an electronic interactive key and to distribution maps are also included.


Introduction
Agathidinae is a moderately large subfamily of Braconidae with 1,069 described species worldwide and 246 in the Oriental Region (Yu et al. 2005), though there are an estimated 2,000-3,000 species awaiting description worldwide (Sharkey et al. 2006).The subfamily has a worldwide distribution and members are found in most terrestrial habitats.The history of higher classification of the Agathidinae was summarized by Sharkey (1992) who also proposed a tribal level classification based on ground-plan coding.Sharkey et al. (2006) conducted phylogenetic analyses based on morphology and the D2-D3 regions of 28S rDNA.The Oriental fauna of Agathidinae were first revised by Bhat and Gupta (1977) and they provided a detailed history of taxonomic research for the area.Sharkey et al. (2009) revised the Oriental genera of Agathidinae.Van Achterberg and Long (2010) revised the Vietnamese agathidine fauna.Sharkey and Clutts (2011) revised the Thai species of Biroia, Braunsia, Camptothlipsis, Coccygidium, Cremnops, Cremnoptoides, Disophrys, Earinus, Gyrochus, Lytopylus, and Troticus, and included a key to the Agathidinae genera of the Oriental region.Sharkey and Stoelb (in press) revised the Thai species of Zelodia.This paper is the third in a series to revise all Thai species of Agathidinae.The genera Aneurobracon, Bassus s.s., Euagathis, and Therophilus s.l. will be dealt with in subsequent publications.

Methods
As part of the inventory of Thai insects, we ran 3 Malaise traps at each of 30 different localities throughout Thailand from 2007-2010, comprising approximately 90 Malaise traps.The specimens dealt with here are primarily from these traps.
Species concepts are based on morphological data and 28S rDNA data.Regions D2-D3 of 28S rDNA (roughly 560 base pairs) were sequenced using the following primers: 28SD2hymF 5' -AGAGAGAGTTCAAGAGTACGTG -3' and 28SD-3hymR 5' -TAGTTCACCATCTTTCGGGTC -3'.Sequences were edited using Geneious Pro v4.7.5 (Drummond et al. 2009) and aligned based on a secondary structure model for Ichneumonoidea developed by Yoder and Gillespie (2004) and Gillespie et al. (2005).Regions of expansion and contraction (RECs), regions of slipped-strand compensation (RSCs), and short regions of alignment ambiguity were further aligned/corrected by eye.Three of these regions (~30 base pairs total) were deleted because they could not be aligned with any confidence, i.e., there were multiple equally supported alignment options.
Phenetic and phylogenetic trees were constructed using neighbor-joining (NJ), maximum parsimony (MP) and Bayesian methods.MP was performed using TNT (Goloboff et al. 2008) [traditional search with 100 random addition sequences followed by branch-swapping, saving 100 trees per replication; 1000 bootstrap replications were used to estimate branch reliability].The Bayesian analysis was performed using MrBayes v3.1.2(Ronquist and Huelsenbeck 2003).Best-fitting DNA substitution models were determined using MrModeltest2.2(Nylander 2004).The general time reversible model of evolution with a parameter for invariant sites and rate heterogeneity modelled under a gamma distribution (GTR+I+G) was determined as the best-fitting model.The Bayesian analysis consisted of two independent Bayesian MCMC runs initiated from different random starting trees.The analysis ran for 5,500,000 generations, reaching a topological similarity criterion of 0.01; trees were sampled every 500 generations.25% of the trees from each run were removed as burn-in upon topological convergence.The NJ tree was produced from PAUP* (Swofford 2003) using default settings.Figure 2 presents the NJ tree, which was more resolved than the phylogenetic trees produced by MP and Bayesian analyses.We mapped the support values of the Bayesian and MP analyses on the NJ tree.
Morphological terms follow Sharkey and Wharton (1997) except for the following: measurements are given for the length and apical width of the first metasomal mediotergite (MT1).Measurement of the apical width is straightforward, however since the base of the tergite is usually hidden from view it is difficult to measure the total length.Instead we measure from the apex of the large tendon that emanates from the propodeum and inserts near the base of the median tergite.Abbreviations used in text: MT1, MT2, MT3...MT7: metasomal mediotergite 1, 2, 3...7.All 14 species found in Thailand are treated with a diagnosis and distributional data.They are illustrated with color photos using a JVC digital camera mounted on a Leica MZ16 microscope and Automontage® stacking software.Distributional data are listed for all species and a Google map via Berkeley Mapper is included for all species.The descriptions are of the holotype and variation is given in parentheses.
The source files for the keys, descriptions, illustrations, DNA sequence and distributional data are all freely available to future researchers who may wish to build on these data.Distribution data, pdf's of non-copywrite references, images, notes, and host and type information can be found by searching Taxabank (a combined specimen and taxonomic database; http://purl.org/taxabank).Codes beginning with an "H" and followed by numbers are unique identifiers used for specimens in the Sharkey lab at the University of Kentucky, and in the specimen database TaxaBank (e.g., H235).All sequences have been deposited in the GenBank database.
Abbreviations used for specimen depositories are as follows:

HIC
Hymenoptera Institute Collection, University of Kentucky, Department of Entomology, Lexington, Kentucky, USA.QSBG Queen Sirikit Botanic Gardens, Chiang Mai, Thailand.

Phylogeny
Recently the polyphyletic generic concept, Bassus, was divided into four genera, i.e., Bassus s.s., a small monophyletic group confined to the Old World, Lytopylus, a large monophyletic group with a world-wide distribution, Neothlipsis, a small New World genus and Therophilus, a polyphyletic dumping ground for the remaining species.Here we refine the concept of Therophilus so that it is monophyletic.However, this leaves those species that do not fit the concept without a correct generic placement.We refer to these as Therophilus s.l. in this treatment, and they will be treated in a separate publication (Sharkey et al. in prep).
The phylogenetic tree in figure 2 shows that Therophilus in the strict sense, and two new genera, are far removed from the majority of the species of Therophilus sensu latu.The clade containing Therophilus s.s. was referred to as an unnamed new tribe in Sharkey (1996).The monophyly of all agathidine tribes is well supported, though the tribal interrelationships based on 28S remain volatile.
Within Therophilus s.s., there are two distinct clades that may deserve generic status when more is known of their biology and diversity.All members of the lower clade, in figure 2, which includes T. cattienensis, are stout and mostly yellow-orange, e.g., all have predominantly yellow-orange heads.Therophilus conspicuous Wesmael, the type species, belongs in this group.Members of the clade containing T. chiangmaiensis are more gracile and melanic.

Type species. Microdus (Therophilus) conspicuous [Lectotype ♀ IRSNB, examined]
Diagnosis.There is neither one character nor a specific combination of characters that distinguishes members of Therophilus from all other agathidines.It is easily separated from members of Cremnoptini and Disophrini by the shape of the tarsal claws, i.e., simple with a basal lobe in Therophilus, versus cleft claws in members of the two aforementioned tribes.Most members of Therophilus have the following combination of diagnostic characters: Apical abscissa of R of fore wing weak at midlength and bent towards apex of wing; interantennal space with two (usually weak and short) longitudinal ridges or with a median keel; postscutellar depression present; cubitus of hind wing strong and tubular, at least basally; subbasal cell of hind wing angled apically at the point from which the cubitus emanates; sclerite between hind coxal cavities and metasomal foramen incomplete or narrow.A combination of characters that distinguishes almost all species of Oriental Therophilus is: Interantennal space with longitudinal groove (Figs.3c, 6c), sometimes weak (Fig. 8c); posteroscutellar depression present (Fig. 8f); notauli sculptured with pits (Fig. 3f).
Legs.Fore tibia lacking pegs, fore tarsal claws with basal lobe; mid tibia with apical and medial pegs; hind tibia with apical pegs.
Mesosoma.Number of pegs on mid tibia = 4. Number of pegs on hind tibia = 9.Sclerite between metasoma and hind coxa wide with a high ridge along most or all of its length.Basal lobe of hind tarsal claw longer than high, not sharply declivous.Lengthwidth of hind femur 1.0/0.342= 2.9.2nd submarginal cell large, cell height subequal to petiole length.Hind wing vein Cub emanates from near mid length of apical margin of subbasal cell, Cub long and strong.Point of notauli intersection heavily sculptured over a wide area with a median longitudinal ridge.Median lobe of mesoscutum bulging and sharply declivous anteriorly.Metapleuron with dense mat of white setae.
Color.Mostly black or dark melanic except for dense white pilosity on metapleuron and white on anterior lateral tergites and sternites, more apical leg segments lighter, tending towards light brown or dark yellow, all apical spurs white, mandible and palpi mostly yellow, fore wing lightly infuscate, stigma brown with a small pale patch near base.Tegula black, concolorous with mesoscutum.Ocellar triangle melanic, concolorous with remainder of vertex.Hind tibia entirely melanic.MT2 entirely melanic.
Etymology.Named in honor of Mr. Anuchat Chaimuangchuen, collector for the TIGER project at Huay Namdung National Park.
Mesosoma.Number of pegs on mid tibia = 5.Number of pegs on hind tibia = 9.Sclerite between metasoma and hind coxa narrow, and lacking a high ridge along its length.Length-width of hind femur 0.596/0.24= 2.5.2nd submarginal cell reduced to a small dot, petiole longer than cell is high, or large, cell height subequal to petiole length.Hind wing vein Cub emanates from near mid length of apical margin of subbasal cell, Cub short and weak.Point of notauli intersection heavily sculptured over a wide area.Median lobe of mesoscutum not bulging and not sharply declivous anteriorly.Metapleuron with scattered white setae.
Metasoma.MT1 length distinctly longer than apical width.MT1 with narrowlyspaced longitudinal striae, with some microsculpture between striae, and lacking two pairs of distinctly stronger striae (carinae).MT1 distinctly wider apically than basally.Ratio of widest point of MT1 to narrowest point 0.32/0.22= 1.5.Length-width ratio of MT1 0.44/0.32= 1.4.MT2 smooth in most of anterior half anteriad transverse groove, longitudinally striate in transverse groove and area posteriad transverse groove, at least medially.Ovipositor much longer than metasoma, about as long as body or longer.
Color.Body mostly melanic, legs mostly pale; body black except as follows: antenna brown, palpi, labrum and other mouthparts yellow, tegula yellow, fore and mid legs entirely yellow, hind coxa mostly black, hind femur mostly brown, paler apically, hind trochanter, tibia, and tarsus mostly yellow, metasomal mediotergite yellow in an- terior half or more, anterior metasomal laterotergites and sternites pale yellow.Tegula yellow, contrasting with predominantly black mesoscutum.Ocellar triangle melanic, concolorous with remainder of vertex.Hind tibia mostly pale, melanic apically and with a subbasal melanic band or lateral spot, or mostly pale, melanic apically only.MT2 pale in anterior half, melanic posteriorly.
Etymology.Named in honor of Mr. Apichat Watanawanit, collector for the TI-GER project at Doi Chiangdao Wildlife Sanctuary.
Description.Body length.4.0 mm.Head.Space between antennal insertions with a weakly developed bulge that is weakly declivous posteriorly, dorsal surface of bulge with a shallow longitudinal groove.Number of flagellomeres 28.Posterior surface of scutellum rugose over a semicircular area that represents the scutellar depression.
Mesosoma.Number of pegs on mid tibia = 5.Number of pegs on hind tibia = 9.Sclerite between metasoma and hind coxa wide with a high ridge along most or all of its length.Length-width of hind femur 0.815/0.256= 3.2.2nd submarginal cell reduced to a small dot, petiole longer than cell is high, or large, cell height subequal to petiole length.Hind wing vein Cub emanates from near anterior apex of apical margin of subbasal cell, Cub short and weak.Point of notauli intersection heavily sculptured over a wide area.Median lobe of mesoscutum not bulging and not sharply declivous anteriorly.
Metasoma.MT1 length distinctly longer than apical width.MT1 with narrowlyspaced longitudinal striae, with some microsculpture between striae, and lacking two pairs of distinctly stronger striae (carinae).MT1 distinctly wider apically than basally.Ratio of widest point of MT1 to narrowest point 0.414/0.278= 1.5.Length-width ratio of MT1 0.668/0.414= 1.6.MT2 smooth in most of anterior half anteriad transverse groove, longitudinally striate in transverse groove and area posteriad transverse groove, at least medially.Ovipositor much longer than metasoma, about as long as body or longer.
Color.Body mostly melanic, legs mostly pale; body black except as follows: antenna brown, palpi, labrum and other mouthparts yellow; tegula yellow; fore and mid legs yellow except for mostly melanic coxae; hind coxa, trochanter, and femur black; hind tibia mostly pale yellow, melanic apically and with a very weak patch of light brown sub-basally; hind basitarsomere mostly yellow, remaining tarsomeres mostly melanic; anterior metasomal laterotergites and sternites pale yellow; fore wing weakly infuscate.Scape entirely melanic.Tegula yellow, contrasting with predominantly black mesoscutum.Ocellar triangle melanic, concolorous with remainder of vertex.Hind tibia mostly pale, melanic apically and with a subbasal melanic band or lateral spot, or mostly pale, melanic apically only.Fore tarsus mostly or entirely pale.Pronotum entirely melanic.MT2 entirely melanic.
Etymology.Named in honor of Ms. Yuwadee Areeluck, collector for the TIGER project at Doi Inthanon National Park.
Description.Body length.3.3 mm.Head.Space between antennal insertions with a weakly developed bulge that is weakly declivous posteriorly, dorsal surface of bulge with a shallow longitudinal groove.Number of flagellomeres 28.Posterior surface of scutellum posterior scutellar depression represented by two pits.
Mesosoma.Number of pegs on mid tibia = 5.Number of pegs on hind tibia = 9.Sclerite between metasoma and hind coxa narrow, and lacking a high ridge along its length.Length-width of hind femur 0.755/0.31= 2.4.2nd submarginal cell large, cell height subequal to petiole length.Hind wing vein Cub emanates from near anterior apex of apical margin of subbasal cell, Cub long and weak.Notauli meeting but sculpture not extending outside of well-defined grooves.
Color.Mostly yellow or yellow-orange with some brown and black; head yellow or orange except antenna, vertex, and occiput brown; thorax yellow or orange and brown, darker near crenulae and sutures; propodeum mostly dark brown; fore and mid legs yellow; hind leg mostly yellow except most of tibia and tarsus melanic, hind tibial spurs yellow; metasoma mostly yellow; posterior terga mostly brown, MT1 with some weak melanic color on longitudinal striae; fore wing weakly infuscate.Tegula yellow, contrasting with predominantly black mesoscutum, or yellow or orange, similar in color to predominantly orange or yellow mesoscutum.Ocellar triangle melanic, contrasting with remainder of vertex.Hind tibia mostly melanic, pale color, if present, restricted to extreme base.MT2 entirely or almost entirely pale.Etymology.Named in honor of Mr. Tawatchai Boontham, collector for the TI-GER project at Huay Namdung National Park.
Description.Body length.4.6 mm.Head.Space between antennal insertions with a weakly developed bulge that is weakly declivous posteriorly, dorsal surface of bulge with a shallow longitudinal groove.Number of flagellomeres 28.Posterior surface of scutellum posterior scutellar depression represented by a deep sculptured semicircular pit.
Mesosoma.Number of pegs on mid tibia = 5.Number of pegs on hind tibia = 9.Sclerite between metasoma and hind coxa narrow with a high ridge along most or all of its length.Length-width of hind femur 1.0/0.351= 2.9.2nd submarginal cell large, cell height subequal to petiole length.Hind wing vein Cub emanates from near anterior apex of apical margin of subbasal cell, Cub long and weak.Point of notauli intersection heavily sculptured over a wide area with a median longitudinal ridge.Metapleuron with white setae of moderate density.
Color.Melanic except as follows: mouthparts mostly yellow; mid and hind tibial spurs yellow; hind tibia with a yellow patch medially at mid length; anterior metasomal laterotergites and sternites mostly pale yellow; fore wing infuscate, more so than other species in this revision.Tegula black, concolorous with mesoscutum.Ocellar triangle melanic, concolorous with remainder of vertex.Hind tibia mostly melanic with pale coloration at midlength at least medially.MT2 entirely melanic.
Etymology.Named after the province in which the type specimen was collected.Molecular data.H1853, GenBank Accession: JQ929190.Distribution.Distribution map can be found at http://purl.org/thaimaps/chiangmaiensis.Diagnosis.Ocellar triangle pale, concolorous with remainder of vertex.Scape at least partly pale, especially anteriorly.
Comments.The Thai specimens differ from the holotype only in the color of the metapleuron which is yellow-brown in the type and melanic in all Thai specimens.This same variation is found in Vietnamese males described by van Achterberg and Long (2010).
Comments.The Thai specimen has a slightly longer ovipositor, otherwise very similar to type.
Description.Body length.3.8 mm.Head.Space between antennal insertions with a weakly developed bulge that is weakly declivous posteriorly, dorsal surface of bulge with a shallow longitudinal groove.Number of flagellomeres 29.Posterior surface of scutellum posterior scutellar depression represented by several (2-3) pits.
Mesosoma.Number of pegs on mid tibia = 5.Number of pegs on hind tibia = 8.Sclerite between metasoma and hind coxa narrow with a high ridge along most or all of its length.Length-width of hind femur 0.911/0.292= 3.1.2nd submarginal cell reduced to a small dot, petiole longer than cell is high.Hind wing vein Cub emanates from near anterior apex of apical margin of subbasal cell, Cub short and weak.Notauli meeting but sculpture not extending much outside of welldefined grooves.
Metasoma.MT1 length distinctly longer than apical width.MT1 with narrowlyspaced, longitudinal striae, with some microsculpture between striae, and with and two pairs of slightly stronger striae (carinae).MT1 distinctly wider apically than ba- Color.Mostly melanic except as follows: mouthparts yellow except galea melanic; fore and mid legs yellow except coxae, trochanters and base of femora melanic; hind tibia mostly pale, melanic in apical third with a pale, tan, lateral, spot near base; anterior metasomal laterotergites and sternites mostly pale yellow; fore wing weakly infuscate.Tegula black, concolorous with mesoscutum.Ocellar triangle melanic, concolorous with remainder of vertex.Hind tibia mostly pale, melanic apically and with a subbasal melanic band or lateral spot, or mostly pale, melanic apically only.Fore tarsus mostly or entirely pale.MT2 entirely melanic.

Diagnosis.
Ovipositor clearly shorter than body, about as long as Metasoma.Hind tibia mostly pale, melanic apically and with a subbasal melanic band or lateral spot.
Description.Body length.3.5 mm.Head.Space between antennal insertions with a weakly developed bulge that is weakly declivous posteriorly, dorsal surface of bulge with a shallow longitudinal groove.Number of flagellomeres 29.Posterior surface of scutellum posterior scutellar depression represented by two pits.
Mesosoma.Number of pegs on mid tibia = 6.Number of pegs on hind tibia = 10.Sclerite between metasoma and hind coxa narrow, and lacking a high ridge along its length.Length-width of hind femur 0.750/0.265= 2.8.2nd submarginal cell large, cell height subequal to petiole length.Hind wing vein Cub emanates from near anterior apex of apical margin of subbasal cell, Cub long and weak.Notauli barely meeting and sculpture not extending past meeting point.
Metasoma.MT1 length distinctly longer than apical width.MT1 with narrowlyspaced longitudinal striae, with some microsculpture between striae, and lacking two pairs of distinctly stronger striae (carinae).MT1 distinctly wider apically than basally.Ratio of widest point of MT1 to narrowest point 0.367/0.264= 1.5.Length-width ratio of MT1 0.54/0.387= 1.4.MT2 with transverse and/or diagonal striae in and/or near transverse depression.Ovipositor much longer than metasoma, about as long as body or longer.
Color.Yellow or yellow-orange except as follows: antenna melanic; thorax with melanic infusions along crenulae and sutures; propodeum mostly melanic; MT3-MT7 mostly brown; Wings hyaline.Scape entirely melanic.Tegula yellow or orange, similar in color to predominantly orange or yellow mesoscutum.Ocellar triangle pale, concolorous with remainder of vertex.Hind tibia mostly pale, melanic apically only.MT2 entirely or almost entirely pale.
Etymology.Named in honor of Mr. Songran Chaksu, collector for the TIGER project at Doi Chiangdao Wildlife Sanctuary.
Description.Body length.3.9 mm.Head.Space between antennal insertions with a weakly developed bulge that is weakly declivous posteriorly, dorsal surface of bulge with a shallow longitudinal groove.Number of flagellomeres 27.Posterior surface of scutellum posterior scutellar depression represented by two pits.
Mesosoma.Number of pegs on mid tibia = 4. Number of pegs on hind tibia = 9.Sclerite between metasoma and hind coxa narrow with a high ridge along most or all of its length.Length-width of hind femur 0.8/0.266= 3.0.2nd submarginal cell large, cell height subequal to petiole length.Hind wing vein Cub emanates from near anterior apex of apical margin of subbasal cell, Cub long and weak.Point of notauli intersection heavily sculptured over a wide area.
Metasoma.MT1 length distinctly longer than apical width.MT1 with narrowlyspaced longitudinal striae, with some microsculpture between striae, and lacking one pair of distinctly stronger striae (carinae).MT1 distinctly wider apically than basally.Ratio of widest point of MT1 to narrowest point 0.383/0.263= 1.5.Length-width ratio of MT1 0.61/0.383= 1.6.MT2 smooth in most of anterior half anteriad transverse groove, longitudinally striate in transverse groove and area posteriad transverse groove, at least medially.Ovipositor much longer than metasoma, about as long as body or longer.
Color.Body mostly melanic, legs mostly pale; body melanic except as follows: mouthparts yellow, tegula brown, fore and mid legs mostly yellow except coxae melanic, basal half of femora brown and apex of tibiae and some tarsomeres light brown, hind tibia mostly pale except apex and a light brown subbasal lateral spot, all tibial spurs yellow; anterior metasomal laterotergites and sternites pale yellow; fore wing weakly infuscate.Tegula black, concolorous with mesoscutum.Ocellar triangle melanic, concolorous with remainder of vertex.Hind tibia mostly pale, melanic apically and with a subbasal melanic band or lateral spot.Pronotum entirely melanic.MT2 entirely melanic.
Etymology.Named in honor of Ms. Acharaporn Sukpeng collector for the TIGER project at Chae Son National Park.
Description.Body length.3.8 mm.Head.Space between antennal insertions with a weakly developed bulge that is weakly declivous posteriorly, dorsal surface of bulge with a shallow longitudinal groove.Number of flagellomeres 28.Posterior surface of scutellum posterior scutellar depression represented by several pits forming a semicircular area.
Mesosoma.Number of pegs on mid tibia = 6.Number of pegs on hind tibia = 8.Sclerite between metasoma and hind coxa narrow with a high ridge along most or all of its length.Length-width of hind femur 0.823/0.256= 3.2.2nd submarginal cell large, cell height subequal to petiole length.Hind wing vein Cub emanates from near anterior apex of apical margin of subbasal cell, Cub short and weak.Notauli extending past meeting point but sculpture not extending over a wide area.
Etymology.Named in honor of Mr. Charoen Wanna, collector for the TIGER project at Doi Phuka National Park.lobes of mesoscutum.Hind tibia largely pale, melanic apically and with a subbasal melanic band or lateral spot, or mostly pale, melanic apically only.
Description.Body length.3.5 mm.Head.Space between antennal insertions with a weakly developed bulge that is weakly declivous posteriorly, dorsal surface of bulge with a shallow longitudinal groove.Number of flagellomeres 29.Posterior surface of scutellum posterior scutellar depression represented by two pits.
Mesosoma.Number of pegs on mid tibia = 5.Number of pegs on hind tibia = 9.Sclerite between metasoma and hind coxa narrow, and lacking a high ridge along its length.Length-width of hind femur 0.850/0.298= 2.9.2nd submarginal cell large, cell height subequal to petiole length.Hind wing vein Cub emanates from near anterior apex of apical margin of subbasal cell, Cub long and weak.Notauli extending past meeting point but sculpture not extending over a wide area.
Metasoma.MT1 length distinctly longer than apical width.MT1 with narrowlyspaced longitudinal striae, with some microsculpture between striae, and lacking two pairs of distinctly stronger striae (carinae).MT1 not distinctly wider apically than basally.Ratio of widest point of MT1 to narrowest point 0.4/0.311= 1.3.Length-width ratio of MT1 0.642/0.4= 1.6.MT2 with transverse and/or diagonal striae in and/or near transverse depression.Ovipositor much longer than metasoma, about as long as body or longer.
Color.Orange, yellow, black, and brown; head mostly orange, ocellar triangle melanic; antenna melanic; mesoscutum mostly pale medially, melanic laterally; prothorax yellow; meso and metapleuron and propodeum melanic; fore and mid legs yellow; hind leg brown except trochanter and most of tibia yellow; metasomal mediotergites mostly melanic except base of MT1 and all of MT2 yellow; anterior metasomal laterotergites and sternites pale yellow, remainder of metasoma melanic; fore wing weakly infuscate.Scape entirely melanic.Tegula yellow, contrasting with predominantly black mesoscutum.Ocellar triangle melanic, contrasting with remainder of vertex, or pale, concolorous with remainder of vertex.Hind tibia mostly pale, melanic apically and with a subbasal melanic band or lateral spot, or mostly pale, melanic apically only.Fore tarsus mostly or entirely pale.MT2 entirely melanic, or pale in anterior half, melanic posteriorly, or entirely or almost entirely pale.
Etymology.Named in honor of Mr. Prasit Wongchai, collector for the TIGER project at Doi Phahompok National Park.
Mesosoma.Number of pegs on mid tibia = 8.Number of pegs on hind tibia = 11.Sclerite between metasoma and hind coxa narrow with a high ridge along most or all of its length.Length-width of hind femur 0.854/0.263= 3.3.2nd submarginal cell reduced to a small dot, petiole longer than cell is high.Hind wing vein Cub emanates from near mid length of apical margin of subbasal cell, Cub long and strong.Notauli extending past meeting point but sculpture not extending over a wide area.Metasoma.MT1 length distinctly longer than apical width.MT1 with narrowlyspaced longitudinal striae, with some microsculpture between striae, and lacking two pairs of distinctly stronger striae (carinae).MT1 distinctly wider apically than basally.Ratio of widest point of MT1 to narrowest point 0.533/0.290= 1.8.Length-width ratio of MT1 0.717/0.533= 1.4.MT2 with short longitudinal striae restricted to transverse depression.Ovipositor much longer than metasoma, about as long as body or longer.Ovipositor length exposed portion of ovipositor distinctly longer than body.
Etymology.Named in honor of Mr. Nikom Wongwan, collector for the TIGER project at Doi Phuka National Park.

Figure 1 .
Figure 1.Map showing Therophilus collection sites in Thailand.

Figure 2 .
Figure 2. NJ phylogram based on 28S rDNA.Where Bayesian and parsimony analyses agreed with the NJ tree, branch support values are included in the figure, i.e., Bayesian posterior probabilities/parsimony bootstrap (values below 0.5 posterior probability and/or 50% bootstrap support were not recorded on the tree).Therophilus s.s.highlighted in grey box.

Figure 3 .
Figure 3. Therophilus anuchati sp.n. a lateral habitus b Wings c dorsolateral head d anterodorsal head e lateral mesosoma f dorsal head and mesosoma g dorsal propodeum h dorsal metasoma.

Figure 4 .
Figure 4. Therophilus apichati sp.n. a lateral habitus b Wings c lateral head and mesosoma d dorsolateral head e dorsal head, mesosoma and propodeum f dorsal Metasoma.

Figure 5 .
Figure 5. Therophilus areeluckae sp.n. a lateral habitus b Wings c anterodorsal head d dorsal head e lateral head and mesosoma f dorsal head and mesosoma g dorsal propodeum and Metasoma.

Figure 6 .
Figure 6.Therophilus boonthami sp.n. a lateral habitus b Wings c anterodorsal head d dorsal head e lateral head and mesosoma f dorsal habitus.

Figure 7 .
Figure 7. Therophilus chiangmaiensis sp.n. a lateral habitus b Wings c anterodorsal head d dorsal head e lateral head and mesosoma f dorsal head and mesosoma g dorsal propodeum h dorsal Metasoma.

Figure 10 .
Figure 10.Therophilus kwanuiae n. sp. a lateral habitus b wings c dorsolateral head d dorsal head e lateral head and mesosoma f dorsal head and mesosoma g dorsal propodeum and metasomal terga 1-3.

Figure 11 .
Figure 11.Therophilus planifrons van Achterberg & Long.a lateral habitus b Wings c dorsolateral head d dorsal head e lateral head and mesosoma f dorsal head and mesosoma g dorsal propodeum and MT1-MT3.

Figure 12 .
Figure 12.Therophilus songrani sp.n. a lateral habitus b Wings c dorsal head d lateral head and mesosoma e dorsal head and mesosoma f dorsal propodeum g dorsal propodeum and Metasoma.

Figure 13 .
Figure 13.Therophilus sukpengae sp.n. a lateral habitus b Wings c anterior head d lateral head and mesosoma e dorsal head and mesosoma f dorsal propodeum and Metasoma.

Figure 14 .
Figure 14.Therophilus wannai sp.n. a lateral habitus b Wings c anterior head d dorsal head e lateral head and mesosoma f dorsal head and mesosoma g dorsal propodeum h dorsal MT1-MT3.

Figure 15 .
Figure 15.Therophilus wongchaii sp.n. a lateral habitus b Wings c anterodorsal head d later head and mesosoma e dorsal head and mesosoma f dorsal propodeum g dorsal MT1-MT3.

Figure 16 .
Figure 16.Therophilus wongwani sp.n. a lateral habitus b Wings c anterodorsal head d lateral head and mesosoma e dorsal mesosoma and propodeum f dorsal Metasoma.