Ten new genera of Agathidini (Hymenoptera, Braconidae, Agathidinae) from Southeast Asia

Abstract The Agathidini (Braconidae: Agathidinae) genera of Southeast Asia are revised based on a phylogenetic analysis of COI and 28S. Ten new genera are proposed, i.e., Agathigma, Asperagathis, Chimaeragathis, Cymagathis, Liragathis, Leuroagathis, Scabagathis, Trochantagathis, Xanthagathis, and Zosteragathis. An illustrated key to the Southeast Asian genera of this tribe is presented. Species from Thailand are keyed and described for all genera of Agathidini except Bassus and Zosteragathis which have too many species for this publication and will be dealt with separately. The phylogenetic analyses indicate that Bassus s.s. is polyphyletic. However, there are no morphological characters to support this and we have retained the current concept of Bassus, which is basically those Agathidini with simple tarsal claws. Numerous new combinations are proposed based on species that are moved to the newly erected genera.


Introduction
Agathidinae is a moderately diverse subfamily of Braconidae with about 1,200 described species (Yu et al. 2012) and many times that number are yet to be named. Larvae are parasitoids of lepidopteran caterpillars of a multitude of families. Most agathidine genera, and probably all of the genera treated here, attack an early instar caterpillar and are quiescent until the host has reached the final instar and is ready to spin a cocoon. At this point in time the parasitoid larva becomes active and quickly consumes the host, i.e., they are koinobiont endoparasitoids. This is the sixth publication on the Agathidine fauna with a concentration on Thailand. Sharkey et al. (2009) revised the Oriental genera of Agathidinae. Sharkey and Clutts (2011) revised the Thai agathidine genera with one or a few species and updated the generic key to the Oriental genera. Stoelb (2012, 2013) revised the Thai species of Zelodia van Achterberg and Agathacrista Sharkey. Lastly, van Achterberg et al. (2014) revised the Thai species of Euagathis. It is the aim of this paper to revise the genera of Agathidini that have not been treated and to describe the Thai species of these that are not overly species-rich.
The recent redefinition of Bassus (Sharkey et al. 2009) to refer only to those species of Old World agathidines with simple claws, necessitated the erection or resurrection of numerous genera to house species formerly contained in the broader, polyphyletic concept of Bassus Sharkey and Stoelb 2013;Sharkey et al. 2016;Achterberg and Long 2010). The previously published genera of this nature are: Gelastagathis Aphelagathis Sharkey, 2015;Pneumagathis Sharkey, 2015;Agathacrista Sharkey, 2013;Neothlipsis Sharkey, 2011;Gyragathis Achterberg & Long, 2010;Aerophilus Szépligeti,1902;and Therophilus Wesmael, 1837. Most of the aforementioned genera, including Bassus s.s., are small and restricted to the Old or New world. The two exceptions are Aerophilus, and Therophilus. These are both species-rich and cosmopolitan. Unfortunately, but perhaps necessarily, Therophilus has become the new dumping ground for unplaced members of Agathidini (Stevens et al. 2010(Stevens et al. , 2011Achterberg and Long 2010). This is all the worse because most of the species recently placed in Therophilus are not closely related to it. Therophilus is sister to the clade Mesocoelus + Aneurobracon and has a number of unique features as outlined in . It is the purpose of this paper to erect new Old World genera to avoid the further debasement of Therophilus. The revision is primarily based on material collected in Thailand. New species from Thailand are keyed and described for all genera of Agathidini except Bassus and Zosteragathis which have too many species for this publication and will be dealt with separately.

Methods
All specimens except for some duplicates are deposited in the Entomological Museum of The Queen Sirikit Botanic Gardens, Chaing Mai, Thailand.

Diagnoses
Diagnoses are rather comprehensive however an abbreviated diagnosis for each genus is given in bold font within each diagnosis.

Morphological terms
Morphological terms are from Sharkey and Wharton (1997) and are matched to the Hymenoptera Anatomy Ontology (HAO; Yoder et al. 2010; http://portal.hymao.org/projects/32/public/ontology/). Identifiers (URIs) in the format http://purl.obolibrary.org/ obo/HAO_XXXXXXX represent anatomical concepts in HAO version http:// purl.obolibrary.org/obo/hao/2011-05-18/hao.owl. They are provided to enable readers to confirm their understanding of the anatomical structures being referenced. To find out more about a given structure, including images, references and other metadata, use the identifier as a web-link, or use the HAO:XXXXXXX (note colon replaces underscore) as a search term at http://glossary.hymao.org. In this paper, terms are linked to the ontology in the results section, each couplet of the key, and in the first description of a taxon (genus Aphelagathis). From this point forward, only terms that do not appear in these areas are hyperlinked.

DNA extraction, PCR and sequencing
DNA was extracted from individual legs with the QIAGEN DNeasy Blood and Tissue Kit using the animal tissue protocol (QIAGEN Inc., Chatsworth, California, USA). The nuclear 28S, regions D2-D3 (~600 bp), rDNA and mitochondrial COI (~650 bp) genes were amplified with the 28S primer pairs 28SD2F (Belshaw and Quicke 1997) and D3R (Harry et al. 1996) and the COI primer pairs LepF1 and LepR1 (Hebert et al. 2004). For COI, PCR was conducted using Takara reagents, with each reaction consisting of 1X buffer, 0.3 mM nucleotides, 0.4 μM of each primer, 0.625 U Takara Ex Taq, ddH2O, and 1-3 μL template DNA in a total reaction volume of 25 μL. The thermal cycling protocol had an initial denaturation period at 95 °C for 2.5 min, followed by 40 cycling steps which denatured at 95 °C for 30 s, annealed at 44 °C for 30 s and extended at 68 °C for 45 s, with a final extension step of 72 °C for 7 min. For 28S, PCR consisted of Qiagen 1X buffer, 4 mM MgSO4, 0.3 mM dNTP, 0.4 μM of each primer, 1.0 U Qiagen Taq, ddH2O, and 1-3 μL template DNA with a total reaction volume of 25 μL. Thermal cycling was as above except annealing at 53 °C, extending for 70 s, and a total of 35 cycles. To determine reaction success, PCR products were electrophoresed in 1% agarose stained with ethidium bromide. PCR products were outsourced for Sanger sequencing either by the Advanced Genetic Technologies Center (University of Kentucky, Lexington, KY) or Beckman Coulter Genomics (Danvers, MA, USA) using labelled dideoxynucleotides with ABI 3730, Big-Dye Terminator mix v. 3.0 or with ABI PRISM 3730xl, BigDye Terminator mix v. 3.1 (Applied Biosystems, Foster City, California, USA).

DNA assembly and phylogenetic analysis
Bi-directional sequences were aligned and edited using Geneious Pro (v. 6.1.5;Drummond et al. 2009) and multiple alignments were assembled using MAFFT (v. 5;Katoh et al. 2006) using the default settings and refined by eye. Maximum likelihood (ML) phylogenetic analyses were conducted on a concatenated (using MacClade v. 4.08; Maddison and Maddison 2000) 1,313-character total evidence data set (COI = 723 bp, 28S = 590 bp) using Garli (v. 2.01;Zwickl 2006). The data were partitioned by gene region and codon position (COI: 3 partitions; 28S: unpartitioned, total of 4 partitions). We applied the most complex model available (GTR+I+G;Rodriguez et al. 1990) to each partition as per recommendations of Huelsenbeck and Rannala (2004). We conducted a 20-replicate ML search for the tree of highest log-likelihood and a 500-replicate ML bootstrap analysis (Felsenstein 1985). Both analyses used the default settings. The data sets analyzed herein are available from the authors upon request.

Phylogenetic considerations
Here we treat a number of species from Thailand and propose 10 new genera. Most of these are demonstratively monophyletic and morphologically distinct; however, some compromises are made due to poor resolution in the phylogenetic analysis. The tree of highest log-likelihood is presented in Figure 1, with the ML bootstrap values plotted on nodes with ≥50% bootstrap support.
In the case of the Thai fauna treated here, there were a number of options in terms of the number of genera that could be proposed. The criteria that I (MJS) used in making decisions on generic limits were: to recognize those monophyletic clades with high (usually > 90%) ML bootstrap support values ( Fig. 1), which are also clearly diagnosed by morphological character states, and the recognition of which would not render other genera paraphyletic. A secondary criterion was to rely solely on potential morphological synapomorphies when they were not contradicted by molecular evidence, as in the case of Bassus (see below). Most of the genera are well-supported by molecular evidence as demonstrated in Figure 1 and by morphological synapomorphies; e.g., Agathacrista, Asperagathis, Camptothlipsis, Chimaeragathis, Liragathis, Neothlipsis, and Trochantagathis. However, Bassus was polyphyletic forming two clades. The members of clades are not distinct morphologically, and they share the character state of lacking a lobe at the base of the tarsal claws. Rather than dividing Bassus into two indistinguishable genera, we prefer to continue to recognize the current concept. Agathigma, Cymagathis, Gyragathis, Leuroagathis, Scabagathis, and Xanthagathis are each represented by only one species; therefore there are no nodes from which to obtain bootstrap values. However, all are on relatively long branches on the total evidence tree, all have distinct autapomorphies, and none renders another taxon paraphyletic (Fig. 1). Zosteragathis is most likely a paraphyletic genus. Although members have similarities, there is not a morphological or molecular autapomorphy for the group. The ML bootstrap values (plotted on Fig. 1) support multiple monophyletic clades of Zosteragathis, but none of these have an obvious morphological autapomorphy. Rather than propose a number of vague genera, I (MJS) thought it best to propose a conservative hypothesis in the interest of stability.

Discussions of each genus are presented below in alphabetical order
Agathacrista Sharkey, 2013: The genus was described and revised by Sharkey and Stoelb (2013). The thin interantennal crest is an autapomorphy for the genus, though convergently found in Chimaeragathis and in a few species of a few other genera; e.g., Therophilus. Agathigma Sharkey, new genus: Agathigma templei is the sole species. Morphological autapomorphies are the squared temples ( Fig. 2) and the labial palpus reduced to 2 segments. The former character state is rarely found in other agathidine genera such as Gyragathis and convergently in a few New World Therophilus and Aerophilus. In the tree in Figure 1 it appears as sister to the clade Leuroagathis + Xanthagathis; however there is no bootstrap support for this relationship. The branch leading to the terminus A. templei is the longest of all branches, a fact that further erodes confidence in its placement. Asperagathis Sharkey, new genus: This genus is sister to one of the Zosteragathis clades in the total evidence tree (Fig. 1). The rugose sculpture of the mesosoma is a proposed autapomorphy; however, even rougher sculpture occurs in Southeast Asian specimens that author MJS has viewed which may not be congeneric. Bassus Fabricius, 1804: Bassus, the strict definition of which was proposed by Sharkey et al. (2009), is restricted to those Old World agathidines with simple tarsal claws. This autapomorphy is convergently found in all Sesioctonus, a Neotropical genus, and in a few species of other genera such as Neothlipsis. Bassus is polyphyletic in the ML tree (Fig. 1); and the polyphyly is not resolved in the ML bootstrap tree (not shown). Interestingly, both COI and 28S gene-trees (analyzed as above; not shown) are completely congruent with the tree in Figure 1 regarding Bassus, showing the same species membership in the 2 clades. Because there are no obvious morphological character states to distinguish either of the two clades of Bassus, we choose to retain the genus as it is until more data confirm that it is not monophyletic. Camptothlipsis Enderlein, 1920: This is an Old World genus, primarily tropical, that is sister to the New World genus Neothlipsis in Figure 1 (but see Fig. 1 in . Both lack strong sculpture on the metasomal median tergites and possess granulate sculpture on metasomal median tergites 1-3. Chimaeragathis Sharkey, new genus: An interantennal crest is shared convergently with members of Agathacrista. Another autapomorphy is the relatively dense pilosity on the scutellar triangle and the lateral areas of the propodeum. The total evidence tree (Fig. 1) shows a sister-group relationship with ((Gyragathis + Cymagathis) Liragathis) but this relationship lacks bootstrap support. Cymagathis Sharkey, new genus: An autapomorphy for the genus is that the second median tergite is covered with strong smooth striae that end evenly at the apex of the tergite with the striae forming a semicircular pattern anteromedially. This is convergently found in some species of Trochantagathis. It is sister to Gyragathis on the total evidence tree ( Fig. 1), supported with a bootstrap value of 90. Gyragathis Achterberg & Long, 2010: An autapomorphy for the genus is that the antennal sockets are margined with carinae. Other possible autapomorphies include the interantennal space with a longitudinal depression and the squared temples, the latter of which is shared convergently with Agathigma. Possession of margined antennal sockets is a character state shared by several distantly related New World Agathidini genera, e.g., Alabagrus and Trachagathis, as well as some genera of Cremnoptini and Disophrini. Gyragathis is sister to Cymagathis on the total evidence tree ( Fig. 1), supported with a bootstrap value of 90. Leuroagathis Sharkey, new genus: This genus possesses two autapomorphic character states: notauli absent, and median tergite 1 smooth, lacking sculpture. It is sister to Xanthagathis in the total evidence tree but the relationship lacks bootstrap support ( Fig. 1). Many agathidines from Australia share these two autapomorphies. The one Australian specimen with these characteristics for which we obtained 28S and COI data does not fall within the clade examined herein (unpublished). Liragathis Sharkey, new genus: An autapomorphy is the median carina of the first median tergite which is as strong as, or stronger than, the lateral carinae. It is sister to Gyragathis + Cymagathis but this relationship lacks bootstrap support (Fig. 1). Scabagathis Sharkey, new genus: There are two autapomorphic character states. The vertex has rough sculpture and the labial palpus, normally 4-segmented, is 3-segmented, with the third palpomere lacking. The total evidence tree ( Fig. 1) shows this genus arising early in the evolution of this group and is sister to a clade containing all genera except Aerophilus, Alabagrus, and Braunsia. Xanthagathis Sharkey, new genus: It is sister to Leuroagathis in the total evidence tree, but the relationship has a low bootstrap support ( Fig. 1; bootstrap value = 64). The pale coloration (particularly the yellow head) is autapomorphic. Other potential autapomorphic states are the hyaline wings and the smooth second median tergite. Zosteragathis Sharkey, new genus: There are no obvious morphological synapomorphies for Zosteragathis and its monophyly is not supported (Fig. 1). Most species have fine longitudinal striations on the second metasomal median tergite and most have a white transverse band on the same tergite. Neither of these is universal and the striations are found in other genera. Members of Zosteragathis are recovered in five separate clades in the total evidence tree (Fig. 1). Monophyly of the genus is not falsified in the total evidence bootstrap tree (not shown) where seven Zosteragathis clades fall into a large polytomy that includes all genera in the tree except Aerophilus, Alabagrus, and Braunsia. Although monophyly of Zosteragathis is dubious, it seems preferable to the alternative of erecting new genera for weakly supported clades that have little or no morphological or sequence support. Diagnosis. Body except for fore and mid legs black, hind leg entirely black. Fore wing slightly infuscate in distal half. Antennal sockets not margined with carinae. Interantennal space with a flat triangular elevation that narrows to a short ridge posteriorly approaching the median ocellus. Temple squared in dorsal view. Labial palpus reduced to 2 segments; presumably palpomere 3 is one of the two lost palpomeres. Notauli depressed and partly or entirely pitted. Scutellar triangle smooth with weak sparse punctures. Ventral margin of hind coxal cavities situated below dorsal margin of metasomal foramen. Pegs on anterior surface of fore tibia absent. Hind trochantellus lacking longitudinal carinae. Second submarginal cell of fore wing minute, cell about the same diameter as wing veins. First median tergite almost entirely irregularly striate, lateral longitudinal carina prominent. Second median tergite slightly wider than long and entirely smooth with hints of short striae and some very weak coriarious microsculpture.

Key to Thai genera of Agathidini
Distribution and diversity. Known only from the type specimen collected in Mae Wong National Park, Thailand.
Diagnosis. Body length 4.7 mm. Ovipositor length/body length ratio = 1.0. Interantennal space with a flat triangular elevation that narrows to a short ridge posteriorly approaching the median ocellus. Antenna with 30 flagellomeres. Labial palp reduced, 2-segmented. Notauli pitted anteriorly, smooth posteriorly where they converge. Scutellar triangle and its posterior surface unusually smooth. Scutellar groove with 1 longitudinal ridge. Fore tibia lacking spines or pegs; mid tibia with 3 pegs; hind tibia with 5 pegs. Basal lobe of tarsal claws large and right-angled, claw only extending slightly beyond apex of lobe. RS vein of fore wing slightly sinuate. Second submarginal cell minute. Hind tibial spurs melanic. Hind tibia entirely melanic. Cu-a crossvein of hind wing bent at point where it is intersected by vein Cub. Etymology. Asper is Latin for rough; here it is in reference to the rugose sculpture on the thoracic pleura of members of the genus. Feminine.
Diagnosis. Body predominantly black; head including orbits of eyes black; dorsal apex of pronotum pale yellow or yellowish brown; metasomal terga all black. Fore wing slightly infuscate in apical half or entirely clear/hyaline. Antennal sockets not margined with carinae. Interantennal space with a shallow crest; between the crest and the median ocellus there is a triangular depression flanked by weak smooth carinae. Temple rounded in dorsal view. Third labial palpomere not greatly reduced, about 1/2 as long as apical palpomere. Notauli depressed and entirely sculptured. Mesoscutum with more rough sculpture than most genera, especially posteriorly near junction of notauli. Scutellar triangle rugose or with deep sparse punctures. Sternaulus completely sculptured to epicnemium, metapleuron covered in rough sculpture. Ventral margin of hind coxal cavities situated below dorsal margin of metasomal foramen. Spines or pegs on anterior surface of fore tibia present or absent. Hind trochantellus lacking longitudinal carinae. Second submarginal cell of fore wing varying from minute, cell about the same diameter as wing veins, to petiolate with petiole slightly longer than cell diameter. First median tergite mostly to about 1/2 irregularly striate, lateral longitudinal carina prominent. Second median tergite slightly wider than long and entirely smooth with some very weak coriarious microsculpture, varying to almost entirely irregularly longitudinally striate, with striae terminating evenly near apex of tergite.
Distribution and diversity. Thailand, but undoubtedly more widespread. Biology. Unknown. Etymology. Asper is Latin for rough and refers to the sculpture of the second metasomal median tergite. Diagnosis. Body length 7.6 mm; ovipositor length/body length ratio = 1.0. Interantennal space with a flat triangular elevation that narrows to a short ridge posteriorly and then divides into two short carinae that diverge to either side of the median ocellus. Antenna with 38 flagellomeres. 3 rd labial (penultimate) palpomere long, about ½ as long as apical palpomere. Scutellar groove with 3-4 longitudinal ridges. Fore tibia with about 9 thickened spines concolorous with normal setae; mid tibia with 7 pegs; hind tibia with 8 pegs.   Diagnosis. Body length 4.5 mm; ovipositor length/body length ratio = 1.1. Interantennal space with a flat triangular elevation that narrows to a short ridge posteriorly and then divides into two short carinae that diverge to either side of the median ocellus.

Key to the Thai species of
Antenna with 32 flagellomeres. 3 rd labial (penultimate) palpomere long, more than ½ as long as apical palpomere. Scutellar groove with 3 longitudinal ridges. Fore tibia lacking thickened spines; mid tibia with 6 pegs; hind tibia with 10 pegs. Etymology. Chimaera is a mythological Greek monster with a goat's body, lion's head, and serpent's tail. In this case, the name is a reference to the many diagnostic characters of the genus which are a combination of features each of which diagnoses other agathidine genera, e.g., crest between antennae, fore tibia with thickened spines. Feminine.
Diagnosis. Metapleuron, scutellum, and all but median cell of propodeum thickly setose. Head, including orbits of eye, black; mesosoma black; metasoma variable. Fore wing slightly infuscate in apical half or entirely clear/hyaline. Antennal sockets not margined with carinae. Interantennal space with a high crest that is sharply declivous posteriorly; between the crest and the median ocellus there is a triangular depression flanked by weak smooth carinae. Temple rounded in dorsal view. Third labial palpomere small, less than 1/3 length of apical palpomere. Notauli depressed and partly or entirely pitted. Scutellar triangle rugose. Ventral margin of hind coxal cavities situated below dorsal margin of metasomal foramen. Pegs on anterior surface of fore tibia present. Hind trochantellus lacking longitudinal carinae. Second submarginal cell of fore wing varying from minute, cell about the same diameter as wing veins, to petiolate with petiole longer than cell diameter. First median tergite partly or mostly irregularly striate to rugosostriate, otherwise smooth; lateral carina present, sometimes weak; median carina present, sometimes weak. Second median tergite wider than long and smooth or mostly smooth with some irregular striae.
Distribution and diversity. Undescribed species are found in other Southeast Asian countries.
Biology. Unknown.   Etymology. Named after David Lohman, who collected of one of the specimens in the type series and who serviced Malaise traps in Trang Province for many months.
Diagnosis. Body predominantly black, mesosoma all black, metasomal terga all black, head black except posterior orbit of eyes partly orange. Fore wing slightly infuscate in apical half. Antennal sockets not margined with carinae. Interantennal space with a flat triangular elevation that narrows to a short ridge posteriorly approaching the median ocellus. Temple rounded in dorsal view. Third labial palpomere not greatly reduced, more than 1/2 as long as apical palpomere. Notauli depressed and partly or entirely pitted. Scutellar triangle with dense punctures or aciculations. Ventral margin of hind coxal cavities situated below dorsal margin of metasomal foramen. Pegs on anterior surface of fore tibia present. Hind trochantellus lacking longitudinal carinae. Second submarginal cell of fore wing minute, cell about the same diameter as wing veins. First median tergite evenly and completely covered in strong striae, lateral carinae strong but partly obscured by sculpture. Second median tergite wider than long. Second median tergite entirely covered with strong striae that end evenly at apex of tergite; striae forming semicircular pattern anteromedially.
Distribution and diversity. Known only from the type species in Thailand but probably widespread throughout Southeast Asia.

Gyragathis Achterberg & Long, 2010
Diagnosis. Antennal sockets margined, completely or at least laterally and medially, with carinae. Interantennal space with a longitudinal depression bordered by carinae. Temples squared in dorsal view. Third labial palpomere minute, barely visible, much smaller than apical palpomere. Notauli depressed and partly or entirely pitted. Scutellar triangle smooth or rugose. Ventral margin of hind coxal cavities situated below or in line with dorsal margin of metasomal foramen. Pegs on anterior surface of fore tibia absent. Hind trochantellus lacking longitudinal carinae Distribution and diversity. There are four species, all of which are restricted to the Oriental region (Taiwan, Philippines, and Viet Nam, Thailand). The three previously described species may be distinguished most easily from G. leucosoma sp. n. by the extensive pale color (yellow to orange) on their mesonota. Achterberg and Long (2010), described the genus Gyragathis and the new species G. guyi from Viet Nam. They also transferred three species to the new genus, viz. G. angulosa (Bhat & Gupta, 1977) G. parallela (Chou & Sharkey, 1989) and G. daanyuanensis (Chen & Yang, 2006). The species described here G. leucosoma, is strikingly different from other members of the genus in aspects of sculpture, dimensions, and color, and may belong in its own genus. Molecular data for the described species are lacking to confirm or refute this suspicion.
Diagnosis. Head and mesosoma orange and black (head with black in ocellar triangle only); metasomal terga predominantly black with some white. Fore wing slightly infuscate in apical half. Antennal sockets not margined with carinae. Interantennal space with a flat triangular elevation that narrows to a short ridge posteriorly and then divides into two short indistinct carinae that approach the median ocellus. Temple rounded in dorsal view. Third labial palpomere small, less than 1/3 length of apical palpomere. Notauli completely absent. Scutellar triangle smooth with weak sparse punctures. Ventral margin of hind coxal cavities situated below dorsal margin of metasomal foramen. Pegs on anterior surface of fore tibia absent. Hind trochantellus lacking longitudinal carinae. Second submarginal cell of fore wing minute, cell about the same diameter as wing veins. First median tergite smooth, lacking microsculpture and carina. Second median tergite wider than long. Second median tergite smooth.
Distribution and diversity. Known only from the type species from Thailand. The few Australian Agathidini for which we have COI data do not belong here.
Leuroagathis paulbakeri Sharkey, sp. n. http://zoobank.org/5AEB324A-084A-48DF-9FC1-FBDEA27D954A Etymology. Named in honor of Mr. Paul Baker who obtained the highest mark (100%) in the written exam of Ent. 770 in the fall of 2015. Diagnosis. Body length 4.5 mm; ovipositor length/body length ratio = 0.8. Interantennal space with a flat triangular elevation that narrows to a short ridge posteriorly and then divides into two short carinae that approach the median ocellus. Antenna with 29 flagellomeres. Third labial (penultimate) palpomere small but easily visible, much smaller than apical palpomere. Scutellar groove with 6 longitudinal ridges. Fore tibia with 7-8 thickened spines; mid tibia with 9 pegs; hind tibia with 12 pegs. First median tergite produced laterally around spiracles. Second median tergite widened apically. Etymology. Lira is Latin for ridge, as in the ridge made by a plow in the earth; it is a reference to the median longitudinal ridge on the first metasomal median tergite. Feminine.
Diagnosis. Antennal sockets not margined with carinae. Interantennal space with a flat triangular elevation that narrows to a short ridge posteriorly approaching the median ocellus. Temple rounded in dorsal view. Third labial palpomere, about 1/2 length of apical palpomere. Notauli depressed and partly or entirely pitted. Scutellar triangle smooth or rugose. Dorsal margin of hind coxal cavities situated above ventralmost margin of metasomal foramen. Pegs on anterior surface of fore tibia absent. Hind trochantellus lacking longitudinal carinae. Second submarginal cell of fore wing varying from minute, cell about the same diameter as wing veins, to petiolate with petiole longer than cell diameter. First median tergite mostly with irregular striae, lateral and median carinae strong. Second median tergite wider than long. Second median tergite from mostly smooth with weak striae restricted to transverse depression, to almost completely striate; in the two species the mostly smooth anteromedial area has transverse or semicircular rugosities, much weaker but otherwise similar to those of Cymagathis.
Distribution and diversity. Known from India, Indonesia (Java) and Thailand. Biology. L. javana has been reared from Etiella zinckenella (Pyralidae). New combinations. Liragathis relativa (Bhat and Gupta), comb. n. from Baeognatha. Liragathis javana (Bhat and Gupta), comb. n. from Baeognatha. Etymology. Bao is Thai for light and nai is Thai for eye. The name refers to the pale color of the superior orbit of the eye. Diagnosis. Body length 6.0 mm; ovipositor length/body length ratio = 1.0. Antenna with 35 flagellomeres. Third labial (penultimate) palpomere about ½ as long as apical palpomere. Scutellar groove with 3 longitudinal ridges. Propodeum rugose and mostly glabrous. Superior orbit of eye orange, posterior orbit also orange. Mesoscutum mostly punctate.
Distribution. Known only from the specimens collected in Thailand. For a distribution map go to: http://bit.ly/22WZjoH
Diagnosis. Vertex of head with rugose sculpture. Head and mesosoma both black and orange; metasomal terga mostly black; base of first median tergite whitish; basal half of second median tergite whitish yellow. Fore wing hyaline, not more infuscate in distal half. Antennal sockets not margined with carinae. Interantennal space with a flat triangular elevation that narrows to a short ridge posteriorly. Temple rounded in dorsal view. Third labial palpomere absent, palpus 3-segmented. Notauli depressed and partly or entirely pitted. Scutellar triangle rugose. Dorsal margin of hind coxal cavities situated above ventral-most margin of metasomal foramen. Pegs on anterior surface of fore tibia absent. Hind trochantellus lacking longitudinal carinae.   Bhat & Gupta, 1977 Etymology. Trochanter comes from the Greek trochalos meaning running; here it is a reference the pair of ridges on the hind trochantellus which are diagnostic for the genus. Feminine.
Diagnosis. Head (including posterior orbit of eye) and mesosoma black; metasomal tergites black or black and pale yellow. Fore wing hyaline, not infuscate in distal half. Antennal sockets not margined with carinae. Interantennal space with a flat triangular elevation that narrows to a short ridge posteriorly and then divides into two short indistinct carinae that approach the lateral margins of the median ocellus. Temple rounded in dorsal view. 3 rd labial palpomere minute, barely visible, much smaller than apical palpomere. Notauli depressed and partly or entirely pitted. Scutellar triangle rugose. Dorsal margin of hind coxal cavities situated above ventral-most margin of metasomal foramen. Pegs on anterior surface of fore tibia present. Hind trochantellus with pair of longitudinal carinae. Second submarginal cell of fore wing varying from about the same diameter as vein Rs about 3x that diameter. First median tergite usually entirely striate, sometimes partly smooth, especially basally; lateral and medial carinae strong. Second median tergite wider than long and varying from completely and smoothly striate, to mostly smooth with weak smooth striae; semicircular pattern of striae usually present anteromedially.
Distribution and diversity. Known from Vietnam, Thailand and Malaysia, but undoubtedly more widespread in the Oriental Region. Based on the analysis of sequence data presented in Figure 1 there are three species of Trochantagathis from Thailand. The females of these species are very similar whereas the males appear to be quite different from one another. Males of the more melanic species are almost entirely melanic whereas the males of the other species are only slightly more melanic than their female conspecifics. With the limited molecular data at hand, the sexual dimorphism, and the similarity of the females of the three putative species, it is not possible to tell with confidence which species, if any, corresponds with the type of T. marshi. Therefore, we choose not to describe the two or three new species at this time. The specimens from Vietnam placed in Therophilus marshi by Achterberg and Long (2010) need verification. They match well with the type except for minor color differences, but so too do the three Thai species. The images of Figure 17 are of a congeneric (and perhaps conspecific) male and female (specimens H799 and H965). These images present better illustrations of the generic characters discussed above as well as the color sexual dimorphism.
Specimens Etymology. Xantho is Greek for yellow and is a reference to the predominantly yellow color of the known species. Feminine.
Diagnosis. Head yellow, mesosoma and metasoma predominantly yellow, with or without melanic areas. Fore wing hyaline. Antennal sockets not margined with carinae. Interantennal space with a flat triangular elevation, with a weak shallow ridge posteriorly not as elevated as the triangular elevation. Temple rounded in dorsal view. Third labial palpomere minute, barely visible, much smaller than apical palpomere. Notauli depressed and partly or entirely pitted. Scutellar triangle smooth with weak sparse punctures. Dorsal margin of hind coxal cavities situated above ventral-most margin of metasomal foramen. Pegs on anterior surface of fore tibia absent. Hind trochantellus lacking longitudinal carinae. Second submarginal cell of fore wing minute, cell about the same diameter as wing veins. First median tergite entirely, finely, irregularly striate; lateral carina weak. Second median tergite wider than long. Second median tergite smooth.
Distribution and diversity. Viet Nam and Thailand. See below for the distribution of the Thai specimens.

Xanthagathis mellisoma (Achterberg and Long), comb. n.
Therophilus mellisoma Achterberg & Long, 2010 Diagnosis. Body length 3.7 mm; ovipositor length/body length ratio = 0.64. Interantennal space with a flat triangular elevation, with a weak shallow ridge posteriorly not as elevated as the triangular elevation. Antenna with 27 flagellomeres. Third labial  palpomere reduced, barely visible, much smaller than apical palpomere. Notauli pitted throughout. Scutellar groove with 3 longitudinal ridges. Fore tibia lacking spines; mid tibia with 6 pegs; hind tibia with 5 pegs. The Thai specimen in Figure 17 differs from the holotype in the lack of a melanic patch distally on hind femur.
Variation. Color usually entirely xanthic (yellow) except for brown as follows: most wing veins including stigma, antenna, hind tarsus and apex of hind tibia. Some specimens are more melanic with brown color extending to propodeum, most of hind leg and parts of most terga.
Etymology. Zoster is Greek for belt; here it is a reference to the white band present on the second metasomal median tergite of most species. Feminine.
Diagnosis. Fore wing hyaline, not infuscate in distal half. Antennal sockets not margined with carinae. Temple rounded in dorsal view. Notauli depressed and partly or entirely pitted. Dorsal margin of hind coxal cavities situated above ventral-most margin of metasomal foramen. Pegs on anterior surface of fore tibia absent. Hind trochantellus lacking longitudinal carinae. Second submarginal cell of fore wing petiolate, small to minute. First median tergite entirely, finely, irregularly striate; lateral carina weak. Second median tergite more than 2x longer than wide. Second median tergite usually entirely, finely, irregularly, striate with striae ending evenly near apex of tergite, rarely striae partly absent. Some species have reduced striae on second median tergite and are recognized by the lack of apomorphic structures that distinguish other closely related genera, e.g., claws not simple, interantennal space without a sharply declivous keel, first median tergite without prominent lateral carina or medial carina, fore tarsus without spines or pegs.
Distribution and diversity. Australian, Ethiopian, Oceania, Oriental, and eastern Palearctic regions.

New species combinations
Below is a list of all new combinations that I am aware of. Since the limits, and even the monophyly, of Zosteragathis are uncertain the list will undoubtedly change in the future. Distribution. Known only from the specimens collected in Thailand. For a distribution map go to: http://bit.ly/1MPrTqu