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Research Article
Ten new genera of Agathidini (Hymenoptera, Braconidae, Agathidinae) from Southeast Asia
expand article infoMichael J. Sharkey, Eric Chapman
‡ University of Kentucky, Lexington, United States of America
Open Access

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.

Keywords

Taxonomy, systematics, Agathigma, Asperagathis, Chimaeragathis, Cymagathis, Liragathis, Leuroagathis, Scabagathis, Trochantagathis, Xanthagathis, Zosteragathis

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. Sharkey and 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 et al. 2015; Sharkey and Chapman 2015; Sharkey and Stoelb 2013; Sharkey et al. 2016; Achterberg and Long 2010). The previously published genera of this nature are: Gelastagathis Sharkey, 2015; 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, 2011; Achterberg 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 Sharkey and Stoelb (2012). 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 dideoxy-nucleotides 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.

Results

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.

Figure 1.

Tree of highest log-likelihood from 20 ML search reps of the combined COI+28S data set with bootstrap values ≥50% plotted at the nodes.

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 Sharkey and Chapman 2015). 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.

Key to Thai genera of Agathidini

1 A1. Fore and mid claws cleft. A2. Ovipositor variable, often barely exerted or shorter than 1/2 length of metasoma, rarely longer. Disophrini and Cremnoptini
B1. Fore and mid claws simple with or without a basal lobe. B2. Ovipositor longer than 1/2 length of metasoma 2
2 A. Median tergite 1 entirely smooth 20
B. Median tergite 1 mostly granulate or coriarious 21
C. Median tergite 1 with other sculpture, usually striate 3
3(2) A1. Median tergite 3 usually extensively striate in anterior half or more. A2. Straight carina situated above hind coxal cavities (CC) 4
B1. Median tergite 3 not extensively striate, usually smooth, or rarely, weakly coriarious. B2. If carina exists between hind coxal cavities (CC) then it is curved and dipping below dorsal margin of coxal cavities. 5
4(3) A. Adventitious vein (2RS) on r-m crossvein of fore wing absent or indicated only by slight swelling Aerophilus Szépligeti
B. Adventitious vein (2RS) on r-m crossvein of fore wing present & distinct Braunsia Kriechbaumer
5(3) A. Fore and mid tarsal claws with a basal lobe 6
B. Fore and mid tarsal claws simple Bassus Fabricius
6(5) A. Mouthparts long, galea significantly longer than wide; gena often elongate Agathis Latreille
B. Mouthparts short (normal), galea not longer than wide; gena not especially elongate 7
7(6) A. With carina partly or completely surrounding antennal socket Gyragathis Achterberg & Long
B. Lacking carina partly or completely surrounding antennal socket 8
8(7) A. Hind trochantellus with ventral longitudinal carinae Trochantagathis gen. n.
B. Hind trochantellus lacking ventral longitudinal carinae 9
9(8) A. Vertex of head smooth, with weak punctures 10
B. Vertex of head rugosopunctate Scabagathis gen. n.
10(9) A1. RS+M vein of fore wing mostly or entirely absent. A2. Notauli present 11
B1. RS+M vein of fore wing present and complete. B2. Notauli absent Earinus Wesmael
11(10) A. Sharply declivous crest in interantennal space present 12
B. Sharply declivous crest in interantennal space absent 14
12(11) A. Cub vein of hind wing absent, or if present, clear and weak, not tubular, and not contiguous with cu-a (base) 13
B. Cub vein of hind wing present, tubular and pigmented Therophilus anuchati Sharkey
13(12) A. Scutellar triangle smooth with punctures and sparse setae Agathacrista Sharkey
B. Scutellar triangle rugose or with dense aciculations, sometimes obscured with dense setae Chimaeragathis gen. n.
14(11) A. Median tergite 2 mostly striate with striae coming to an abrupt and uniform end at or near apex of tergite 15
B. Median tergite 2 striate or not; if striate, striae not coming to an abrupt and uniform end at apex of tergite 17
15(14) A. Semicircular striae at base of median tergite 2 present Cymagathis gen. n.
B. Semicircular striae at base of median tergite 2 not present 16
16(15) A. Pegs of fore tibia present (concolorous with tibia therefore difficult to see) Asperagathis gen. n.
B. Pegs of fore tibia absent Zosteragathis gen. n.
17(14) With two or more of the following characters: A1. Rs vein of fore wing weak medially and bent: A2. Sclerite separating hind coxal cavities from metasomal foramen narrow or absent. A3. Apex of scutellum (at border with metanotum) with a distinct, often semicircular, depression. A4. Interantennal space often with two small protrusions separated by a depression (use frontal view). A5. Cub vein of hind wing pigmented and tubular where it is attached to subbasal cell and causing an angle in the distal margin of the cell where it is attached Note: often (70%) small and pale in coloration Therophilus Wesmael
With none or at most one of the above character states. Rather the following character states apply: B1. Rs vein of fore wing evenly sclerotized and straight. B2. Sclerite separating hind coxal cavities from metasomal foramen relatively wide. B3. Apex of scutellum (at border with metanotum) smooth or sculptured but lacking deep depression(s). B4. Interantennal space without two small protrusions separated by a depression (use frontal view), rather smooth, or with a median keel that may or may not be pronounced. B5. Cub vein of hind wing absent, OR not attached to basal cell, OR not pigmented, and subbasal cell not angled at point of intersection. Note: often (70%) small and pale in coloration Note: body usually mostly melanic 18
18(17) A. Median tergite 2 partly or entirely white, ivory, or pale yellow Xanthagathis gen. n.
B. Median tergite 2 entirely melanic 19
19(18) A. Temples rounded. AA. Median longitudinal ridge of first median tergite present Liragathis gen. n.
B. Temples squared. BB. Median longitudinal ridge of first median tergite absent Agathigma gen. n.
20(2) A. RS+M vein of fore wing mostly or entirely absent Leuroagathis gen. n.
B. RS+M vein of fore wing present and complete Earinus Wesmael
21(2) A. RS vein of fore wing completely absent Aneurobracon Brues
B. RS vein of fore wing present, though sometimes interrupted at midlength Camptothlipsis Enderlein
Agathidini key, Figure 1.

Agathidini key, Figure 2.

Agathidini key, Figure 3.

Agathidini key, Figure 4.

Agathidini key, Figure 5.

Agathidini key, Figure 6.

Agathidini key, Figure 7.

Agathidini key, Figure 8.

Agathidini key, Figure 9.

Agathidini key, Figure 10.

Agathidini key, Figure 11.

Agathidini key, Figure 12.

Agathidini key, Figure 13.

Agathidini key, Figure 14.

Agathidini key, Figure 15.

Agathidini key, Figure 16.

Agathidini key, Figure 17.

Agathidini key, Figure 18.

Agathidini key, Figure 19.

Agathidini key, Figure 20.

Agathidini key, Figure 21.

Descriptions

Note: The text in bold font in the diagnoses below show a minimum set of character states to distinguish the taxon. The numbers preceded with the letter H are unique identifiers associated with each specimen.

Agathigma Sharkey, gen. n.

Type species

Agathigma templei Sharkey, sp. n.

Etymology

Aga (from Agathis); thigma is Greek for touch, here used as a reference to the reduced 2-segmented palpi. Feminine.

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.

Distribution and diversity

Known only from the type specimen collected in Mae Wong National Park, Thailand.

Agathigma templei Sharkey, sp. n.

Etymology

Named after Jimmy Temple, childhood friend of the first author; the fact that the temples are squared may be coincidental.

Figure 2.

Agathigma templei holotype female: a labial palpus, arrows indicate the two palpomeres b lateral habitus c dorsal head d wings; arrows from top to bottom indicate: RS vein; minute second submarginal cell; angle in cu-a crossvein of hind wing e anterodorsal head, arrows indicate ridge between antennae f lateral head g lateral mesosoma h dorsal mesonotum i dorsal propodeum j dorsal metasoma.

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.

Specimens examined

Holotype 2♀♀ (H415): THAILAND, Kamphaeng Phet, Mae Wong NP Chong Yen, 16°5.212'N, 99°6.576'E, 1306 m, Malaise trap, 20–27.viii.2007, Piluek C. & Inpuang A. leg.

Distribution

Known only from the type specimen collected in Mae Wong National Park, Thailand. For a distribution map go to: http://bit.ly/22WV8JD

Asperagathis Sharkey, gen. n.

Type species

Asperagathis xesta Sharkey, sp. n.

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.

Key to the Thai species of Asperagathis

1 a. Metasomal median tergite 2 mostly or entirely smooth. aa. Second submarginal cell minute, diameter about equal to thickness of surrounding wing veins Asperagathis xesta Sharkey, sp. n.
b. Metasomal median tergite 2 mostly rugosostriate. bb. Second submarginal cell normal (wider than below), diameter much wider than thickness of surrounding wing vein Asperagathis aspera Sharkey, sp. n.
Asperagathis key, Figure 1.

Asperagathis aspera Sharkey, sp. n.

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. 3rd 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.

Figure 3.

Asperagathis aspera, holotype female: a lateral habitus b fore wing c hind wing d dorsal head e lateral head f lateral mesosoma g dorsal thorax h propodeum and metasomal terga 1-2.

Specimens examined

Holotype ♀ (H274): THAILAND, Phetchabun , Thung Salaeng Luang NP, Pine forest; Gang Wang Nam Yen, 16°35.789'N, 100°52.769'E, 732 m, Malaise trap, 15–22.vi.2007, Pongpitak & Sathit leg.

Distribution

Known only from the type specimen collected in Thung Salaeng Luang National Park, Thailand. For a distribution map go to: http://bit.ly/1T5FqXj

Asperagathis xesta Sharkey, sp. n.

Etymology

Xestos is Greek for smooth and refers to the smooth second metasomal median tergite.

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. Antenna with 32 flagellomeres. 3rd 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.

Figure 4.

Asperagathis xesta Holotype female: a lateral habitus b fore wing c dorsal head d lateral head e lateral mesosoma f dorsal thorax g propodeum h metasomal terga 1-3.

Specimens examined

Holotype ♀ (H095): THAILAND, Chaiyaphum, Tat Tone NP, Lawn near Sab Somboon forest unit, 16°0.792'N, 101°58.472'E, 648m, Malaise trap, 26.xi–3.xii.2006, Tawit Jaruphan leg. Paratype ♀ (H1682): Same data as holotype.

Distribution

Known only from the type specimens collected in Tat Tone National Park, Thailand. For a distribution map go to: http://bit.ly/1VPL5H8

Chimaeragathis Sharkey, gen. n.

Type species

Chimaeragathis eurysoma Sharkey, sp. n.

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.

Key to Thai species of Chimaeragathis

1 a. Hind femur yellow; hind tibia mostly yellow Chimaeragathis lohmani Sharkey, sp. n.
b. Hind femur yellow laterally at mid length, black basally and apically, hind tibia black. Chimaeragathis chrysoma Sharkey, sp. n.
c. Hind femur black; hind tibia mostly black with pale patches basally Chimaeragathis eurysoma Sharkey, sp. n.
Chimaeragathis key, Figure 1.

Chimaeragathis chrysoma Sharkey, sp. n.

Etymology

Chrysoma is Greek for an object made of gold and is a reference to the gold colored setae on the mesosoma.

Diagnosis

Body length 6.9 mm; ovipositor length/body length ratio = 0.8. Antenna with 42 flagellomeres. Third labial (penultimate) palpomere about 1/3 as long as apical palpomere. Propleuron convex, lacking distinct bump. Scutellar groove with 3 longitudinal ridges. Fore tibia with 2 pegs; mid tibia with 5 pegs; hind tibia with 4 pegs. Basal lobe of tarsal claws large, right-angled; claw extending slightly beyond apex of lobe.

Figure 5.

Chimaeragathis chrysoma holotype female: a lateral habitus b wings c anterolateral head d lateral head e lateral mesosoma f dorsal thorax g propodeum and metasomal terga 1-3.

Specimens examined

Holotype ♀ (H710): THAILAND, Petchaburi, Kaeng Krachan NP km33/helipad, 12°50.177'N, 99°20.688'E, 735 m, Malaise trap, 18-25.v.2009, Sirichai leg.

Distribution

Known only from the type specimen collected in Kaeng Krachan National Park, Thailand. For a distribution map go to: http://bit.ly/29nOQlL

Chimaeragathis eurysoma Sharkey, sp. n.

Etymology

Eurys is Greek for wide; soma is Greek for body. The species name refers to the wide metasoma of this species.

Diagnosis

Body length 4.8 mm; ovipositor length/body length ratio = 0.7. Antenna with 34 flagellomeres. Third labial (penultimate) palpomere about 1/3 as long as apical palpomere. Propleuron convex, lacking distinct bump. Scutellar groove with 3 longitudinal ridges. Fore tibia with 3 pegs; mid tibia with 4 pegs; hind tibia with 3 pegs. Basal lobe of tarsal claws large, right-angled; claw extending slightly beyond apex of lobe.

Figure 6.

Chimaeragathis eurysoma, female paratypes: a lateral habitus b wings c anterolateral head d lateral head e lateral mesosoma f dorsal thorax g propodeum h metasomal terga 1-3.

Specimens examined

Holotype ♀ (H925): THAILAND, Petchaburi, Kaeng Krachan NP, km33/helipad, 12°50.177'N, 99°20.688'E, 735 m, Malaise trap, 25.i–4.ii.2009, Sirichai leg. Paratypes: THAILAND: ♀ (H321), Prachuab Khiri Khan, Khao Sam Roi Yot NP Khao Look Glang 12°6.414'N, 99°57.292'E, Malaise trap, 28.ix–5.x.2008, Yai Amnad leg. ♀ (H242), Trang, Khao Pu-Khao Ya NP, 7°33.038'N, 99°47.369'E, 75 m, Malaise trap, 28.ii–1.iii.2006 M Sharkey leg. ♀ (H649), Chanthaburi, Khao Khitchakut NP, nature trail/fern, 12°50.55'N, 102°7.3'E, 50 m, Malaise trap, 1–8.v.2009, Suthida Charoenchai leg. ♀ (H335), Chanthaburi, Khao Khitchakut NP, nature trail/Banyan tree, 12°50.54'N, 102°7.31'E, 90 m, Malaise trap, 1–8.v.2009, Suthida Charoenchai leg. ♀ (H045), Trang, nr. nam Tok Ton Prew Kae Chong, MT, 140 m, 7°33.15'N, 99°47.38'E, 28.i–3.ii.2005 D Lohman. ♀ (H069), Trang, nr. nam Tok Ton Prew Kae Chong, MT, 140 m, 7°33.15'N, 99°47.38'E, 4–11.ii.2005 D Lohman.

Distribution

Known only from the type specimens collected in Thailand. For a distribution map go to: http://bit.ly/1WNrlTX

Chimaeragathis lohmani Sharkey, sp. n.

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 length 6.2 mm; ovipositor length/body length ratio = 0.8. Antenna with 39 flagellomeres. Third labial (penultimate) palpomere about 1/3–1/2 as long as apical palpomere. Propleuron convex, lacking distinct bump. Scutellar groove with 3 longitudinal ridges. Fore tibia with 1 peg; mid tibia with 3 pegs; hind tibia with 4 pegs. Basal lobe of tarsal claws large, right-angled; claw extending slightly beyond apex of lobe.

Figure 7.

Chimaeragathis lohmani. a and c–h Holotype female b fore wing of paratype H412 c anterolateral head d lateral head e lateral mesosoma f dorsal thorax g propodeum h metasomal terga 1-3.

Specimens examined

Holotype ♀ (H072), THAILAND, Trang, Ampuh Nayong Khaochong, 7°33.038'N, 99°47.369'E, 75 m, 14–16.ii.2005, Mal. trap D Lohman. Paratypes: THAILAND: ♀ (H077), Trang, Khaochong, 7°33.038'N, 99°47.369'E, 75 m, 13.vi.2005, Mal. trap. ♀ (H412), Surat Thani, Khao Sok NP Klong Morg Unit, 8°53.725'N, 98°39.025'E, 87 m, Malaise trap, 10–17.ii.2009, Pongphan leg. Malaysia: 2♀♀ (H5932, H5935), Perlis, Wang Kelian, 6°40'40.94"N, 100°11'23.94"E, 2008, Sharkey and Norliyana.

Distribution

Known only from the type specimens collected in northern Malaysia and southern Thailand. For a distribution map go to: http://bit.ly/1r7TE3x

Cymagathis Sharkey, gen. n.

Type species

Cymagathis krikoma Sharkey, sp. n.

Etymology

Cymato is Greek for wave; here it is a reference for the uniform, large, wave-like striae on metasomal median tergite 2. Feminine.

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.

Biology

Unknown

Cymagathis krikoma Sharkey, sp. n.

Etymology

Krikoma is Greek for ring and refers to the half ring-shaped carina on median tergite two.

Diagnosis

Body length 6.0 mm; ovipositor length/body length ratio = 0.9. Scutellar groove with 3 longitudinal ridges. Fore tibia with 4 thickened melanic spines; mid tibia with 3 pegs; hind tibia with 4 pegs. Flagellomeres rather pale colored. Posterior orbit of eye orange. Sternaulus deeply sculptured and long. Metapleuron rugose over most of surface.

Figure 8.

Cymagathis krikoma paratype female: a lateral habitus b wings c dorsal head, arrow indicating orange posterior orbit d lateral head e lateral mesosoma f dorsal thorax g propodeum h metasomal terga 1-3, arrow indicating semicircular carina on median tergite 2.

Specimens examined

Holotype ♀ (H276), THAILAND, Chaiyaphum, Tat Tone NP, Water tank at Tat Fah waterfall, 15°56.468'N, 102°5.855'E, 245 m, Malaise trap, 19–26.iii.2007, Tawit Jaruphan & Orawan Budsawong leg. Paratypes: THAILAND: ♀ (H290), Chaiyaphum , Tat Tone NP, Officer house at Tat Fah waterfall, 15°56.461'N, 102°5.955'E, 242 m, Malaise trap, 12–19.iii.2007, Tawit Jaruphan & Orawan Budsawong leg. ♀ (H5924), Chaiyaphum, Tat Tone NP, Forest fire Protection station, 16°0.809'N, 102°1.335'E, 195 m, Malaise trap, 3–9.vi.2006, Tawit Jaruphan & Orawan Budsawong leg. ♀ (H2401), Phetchabun, Nam Nao NP Check point, 16°43.695'N, 101°33.797'E, 921 m, Malaise trap, 5–12.v.2007, Leng Janteab leg. ♀ (H483), Mae Hong Son, Namtok Mae Surin NP, Haad Saen, 19°20.857'N, 97°59.123'E, Malaise trap, 27.iv–4.v.2008, Na-maadkam, leg.

Distribution

Known only from the specimens collected in Thailand but Bassus transtriatus (Bhat and Gupta) from Philippines may belong here. For a distribution map go to: http://bit.ly/1SWUYfQ

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.

New combinations

Gyragathis sabahensis (Bhat and Gupta), comb. n., from Agathis. Contrary to Achterberg and Long (2010) B. daanyuanensis (Chen & Yang, 2006) is a member of Therophilus, Therophilus daanyuanensis comb. n.

Biology

Unknown.

Gyragathis leucosoma Sharkey, sp. n.

Etymology

Leucosoma is Greek for white body. The species name refers to the dense white setae on the metapleuron.

Diagnosis

Body length 6.1 mm; ovipositor length/body length ratio = 0.9. Antenna missing after 28th flagellomere. Third labial (penultimate) palpomere about 1/3 as long as apical palpomere. Propleuron with distinct bump near ventral margin. Scutellar groove with 5 longitudinal ridges. Fore tibia without pegs or thickened spines; mid tibia with 6 pegs; hind tibia with 13 pegs.

Figure 9.

Gyragathis leucosoma, female holotype. a lateral habitus b wings c dorsal head d lateral head e lateral mesosoma f dorsal thorax g propodeum h metasomal terga 1-3.

Biology

Unknown.

Specimens examined

Holotype ♀ (H275), THAILAND, Nakhon Nayok, Khao Yai NP, Lum Ta Kong View Point, 14°25.762'N, 101°23.527'E, 732 m, Malaise trap, 12–19.iv.2007, Wirat Sukho leg.

Distribution

Known only from the type specimen collected in Khao Yai National Park, Thailand. For a distribution map go to: http://bit.ly/1SWVgDh

Leuroagathis Sharkey, gen. n.

Type species

Leuroagathis paulbakeri Sharkey, sp. n.

Etymology

Leuros is Greek for smooth, level, polished and refers to the lack of notauli and smooth metasomal terga. Feminine.

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.

Biology

Unknown.

Leuroagathis paulbakeri Sharkey, sp. n.

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.

Figure 10.

Leuroagathis paulbakeri holotype female: a lateral habitus b wings c dorsal head d lateral head e lateral mesosoma f dorsal thorax g propodeum h metasomal terga 1-3.

Specimens examined

Holotype ♀ (H369), THAILAND, Prachuab Khiri Khan, Khao Sam Roi Yot NP, foot of Khao Taen, 12°8.75'N, 99°57.988'E, 1 m, Malaise trap, 17–24.v.2009, Yai Amnad leg.

Distribution

Known only from the type specimen collected in Khao Sam Roi Yot National Park, Thailand. For a distribution map go to: http://bit.ly/29hEQ95

Liragathis Sharkey, gen. n.

Type species

Liragathis baonai Sharkey, sp. n.

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 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 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.

Key to Thai Liragathis species

1 a. Mesoscutum mostly or entirely black 2
b. Mesoscutum mostly or entirely orange Liragathis javana (Bhat & Gupta)
2 a. Superior orbit of eye, between antennal insertion and eye, orange Liragathis baonai Sharkey, sp. n.
b. Superior orbit of eye, between antennal insertion and eye, black Liragathis damnai Sharkey, sp. n.
Liragathis key, Figure 1.

Liragathis key, Figure 2.

Liragathis baonai Sharkey, sp. n.

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.

Figure 11.

Liragathis baonai paratype female: a lateral habitus b hind tibia and tarsus c wings d dorsal head e lateral head f lateral mesosoma g dorsal thorax h propodeum and metasomal terga 1-3.

Specimens examined

Holotype ♀ (H360), THAILAND, Nakhon Si Thammarat, Namtok Yong NP, behind campground lavatory, 8°10.434'N, 99°44.508'E, 80 m, Malaise trap, 9–16.ix.2008, U-prai leg. Paratypes: ♀ (H282) THAILAND, Trang, Khao Pu-Khao Ya NP, 7°32.534'N, 99°47.856'E, 145 m, Malaise trap, 2–9.xii.2005 M Sharkey leg. MALAYSIA: 2♀♀ (H5928, H16987), Pahang, Kuala Lompat, 1.ix.1999, 3°41'44.27"N, 102°13'25.42"E, Nor Zaneedarwaty leg. ♀ (H16988), Selangor, Kuala Sawit, 3°11'N, 101°37'E, 22.xi.1999, Nor Zaneedarwaty leg.

Distribution

Known only from the specimens collected in Thailand. For a distribution map go to: http://bit.ly/23QN2Ik

Liragathis damnai Sharkey, sp. n.

Etymology

Dam is Thai for black and nai is Thai for eye. The name refers to the black color of the superior orbit of the eye.

Diagnosis

Body length 5.3 mm; ovipositor length/body length ratio = 0.7. Antenna with 33 flagellomeres. Third labial (penultimate) palpomere about ½ as long as apical palpomere. Scutellar groove with 3 longitudinal ridges. Fore tibia lacking pegs; mid tibia with 3 pegs; hind tibia with 6 pegs. Fore tibia lacking pegs; mid tibia with 3 pegs; hind tibia with 6–8 pegs. Propodeum rugose but with discernible large areolae as in some Lytopylus species. Superior orbit of eye black, posterior orbit orange. Mesoscutum mostly rugose.

Figure 12.

Liragathis damnai paratype female: a dorsal head b lateral habitus b hind tibia and tarsus c dorsal thorax d scutellum and propodeum e metasomal terga 1-3 f fore wing g hind wing.

Specimens examined

Holotype ♀ (H468), THAILAND Chiang Mai, Doi Chiang Dao WS, Pha Tang unit, 19°24.978'N, 98°54.886'E, 526 m, Malaise trap, 24–31.iii.2008, Songkran & Apichart leg. Paratypes: THAILAND: ♀ (H999), Lampang Chae Son NP, Youthcamp/meeting hall, 18°49.866'N, 99°28.209'E, 476 m, Malaise trap, 15-22.iii.2008 B Kwannui & A. Sukpeng leg. ♀ (H2416), Lampang Chae Son NP Youthcamp, 18°49.826'N, 99°28.256'E, 455 m, Malaise trap, 1–7.iv.2008 B Kwannui & A. Sukpeng leg. ♀ (H397), Chiang Mai, Queen Sirikit Botanic Garden, 18°52'57.5"N, 98°51'35.5"E, 17–24.ii.2009, MT K Kaewjanta & R. Sawkord leg.

Distribution

Known only from the specimens collected in Thailand. For a distribution map go to: http://bit.ly/22WZjoH

Liragathis javana (Bhat and Gupta)

Baeognatha javana Bhat & Gupta, 1977.

Diagnosis

Body length 6.0 mm; ovipositor length/body length ratio = 0.8. Antenna with 33 flagellomeres. Third labial (penultimate) palpomere about ½ as long as apical palpomere. Scutellar groove with 3 longitudinal ridges. Fore tibia lacking pegs; mid tibia with 4 pegs; hind tibia with 5 pegs. Posterior orbit of eye orange. Mesoscutum, scutellum, pronotum and part of mesopleuron orange. Second median tergite wide, about two times wider than long. Similar to L. relativa (Bhat and Gupta). Second submarginal cell of L. javana much larger.

Figure 13.

Liragathis javana female: a lateral habitus b wings c dorsal head d lateral head e lateral mesosoma f dorsal thorax g propodeum and metasomal terga 1-3 h holotype lateral habitus.

Specimens examined

Holotype ♀, Indonesia, Java, Bogor (=Buitenzorg), 15.ix.1956, ex. Etiella zinckenella M Satarchi, USNM, examined). Other: Indonesia: 5♀♀, 2♂♂ (H16989 - H16992), Central Java, Tepus, 6°49'S, 110°53'E, 7.v.1990, host Etiella sp. [Pyralidae], coll. G.C. Luther (EMEC).Thailand: ♀ (H628), Phetchabun, Nam Nao NP check point, 16°43.687'N, 101°33.754'E, 924 m, Malaise trap, 5–12.v.2007, Noopean Hongyothi leg. ♀ (H458), Phetchabun, Nam Nao NP check point, 16°43.695'N, 101°33.797'E, 921 m, Malaise trap, 28.iv–5.v.2007, Leng Janteab leg. ♀ (H2406), Phetchabun, Nam Nao NP check point 16°43.695'N, 101°33.797'E, 921 m, Malaise trap, 5–12.v.2007, Leng Janteab leg. ♀ (H419), Kanchanaburi, Khuean Srinagarindra NP, Tha Thung-na/Chong Kraborg, 14°29.972'N, 98°53.035'E, 210 m, Malaise trap, 19–26.iii.2009, Boonnam & Phumarin leg. ♀ (H366), Kanchanaburi, Khuean Srinagarindra NP, Tha Thung-na/Chong Kraborg, 14°29.972'N, 98°53.035'E, 210 m, Malaise trap, 26.iii–2.iv.2009, Boonnam & Phumarin leg.

Distribution

Known only from the specimens collected in Thailand and Indonesia. For a distribution map go to: http://bit.ly/2ajVCca

Scabagathis Sharkey, gen. n.

Type species

Scabagathis emilynadeauae Sharkey, sp. n.

Etymology

Scaber is Latin for rough, scabby, mangy; here it refers to the rough (rugose) sculpture on the vertex of the head. Feminine.

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. 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 longer than wide. Second median tergite entirely, finely, irregularly striate, with striae ending evenly near apex of tergite.

Distribution and diversity

Known only from the type species from Thailand.

Biology

Unknown.

Scabagathis emilynadeauae Sharkey, sp. n.

Etymology

Named in honor of Ms. Emily Nadeau who obtained the highest mark in the weekly quizzes of Ent. 770 in the fall of 2015.

Diagnosis

Body length 5.1 mm; ovipositor length/body length ratio = 0.6. Antenna with 31 flagellomeres. Scutellar groove with 3 longitudinal ridges. Fore tibia without thickened spines or pegs; mid tibia with 2 pegs; hind tibia missing. First median tergite whitish at extreme base. Second median tergite whitish in basal half.

Figure 14.

Scabagathis emilynadeauae holotype female: a lateral habitus b wings c dorsal head d lateral head e lateral mesosoma f dorsal thorax g propodeum h metasomal terga 1-3.

Specimens examined

Holotype ♀ (H033), Thailand, Trang, Nayong, Khaochong, 24–27.vi.2005, 7°33.038'N, 99°47.369'E, 75 m, Malaise trap.

Distribution

Known only from the type specimen collected in Thailand. For a distribution map go to: http://bit.ly/29kPFrZ

Trochantagathis Sharkey, gen. n.

Type species

Baeognatha marshi 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. 3rd 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, 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.

Biology

Unknown.

New combinations

Trochantagathis marshi (Bhat and Gupta), comb. n., from Baeognatha.

Trochantagathis marshi (Bhat and Gupta), comb. n.

Baeognatha marshi Bhat & Gupta 1977

Therophilus marshi, Achterberg & Long, 2010

Diagnosis

Body length 5.6 mm; ovipositor length/body length ratio = 0.7. Antennae broken (37–38 flagellomeres in Thai congenerics). Third labial (penultimate) palpomere about 1/5 as long as apical palpomere. Scutellar groove with 3 longitudinal ridges. Fore tibia with 2 pegs; mid tibia with 5 pegs; hind tibia with 5 pegs.

Figure 15.

Trochantagathis marshi holotype female: a dorsal habitus b hind femur showing one of the two ridges on the trochantellus.

Figure 16.

Trochantagathis marshi? female: a lateral habitus b propodeum and metasomal terga 1-2 c fore wing d detail of ridges on trochantellus of hind leg e lateral view of hind leg and metasoma of male (H965); note melanic color of hind coxa and femur.

Specimens examined

Holotype ♀, Malaysia, Sabah, Bettotan nr. Sandakan, 15.viii.1927 (FSCA).

Distribution

Malaysia, Vietnam and Thailand. For a distribution map of the Thai specimens go to: http://bit.ly/1VK7I0a

Xanthagathis Sharkey, gen. n.

Type species

Therophilus mellisoma Achterberg & Long, 2010.

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.

Biology

Unknown.

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.

Figure 17.

Xanthagathis mellisoma, female: a lateral habitus b labial palp, arrow indicates minute third palpomere c wings d dorsal head e lateral head f lateral mesosoma g dorsal thorax h propodeum i metasomal terga 1-3.

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.

Distribution

For a distribution map of the Thai specimens go to: http://bit.ly/1SWVASF

Zosteragathis Sharkey, gen. n.

Type species

Zosteragathis samensis Sharkey, sp. n.

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.

Biology

Hosts are unknown for all Thai species; however, there are records for three extra-Thai species. These appear to suggest that the host range is wide. The records are: Zosteragathis coryphe was reared from Phycodes radiata (Sesioidea: Brachodidae) (Nixon 1950). Zosteragathis robusta (Achterberg and Long) from Vietnam was reared from “Omiodes indicata (Lepidoptera: Pyralidae: Pyraustinae) on soybean (Glycine max (Linnaeus)), according to the label data”, (Achterberg and Long 2010). Zosteragathis festiva (Muesebeck) was reared from Grapholitha molesta, the oriental fruit moth, (Tortricoidea: Tortricidae). Many other Lepidoptera from a wide range of families are listed by Yu et al. (2012) as hosts of Z. festiva, e.g., Blastobasidae, Carposinidae, Gelechiidae, Noctuidae, and Pyralidae.

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.

Zosteragathis annulus (Chou & Sharkey, 1989), comb. n. from Bassus

Zosteragathis asper (Chou & Sharkey, 1989), comb. n. from Bassus

Zosteragathis conformis (Bhat & Gupta, 1977), comb. n. from Agathis

Zosteragathis contrasta (Achterberg & Long, 2010), comb. n. from Therophilus

Zosteragathis coryphe (Nixon, 1950), comb. n. from Agathis

Zosteragathis depressa (Chou & Sharkey, 1989), comb. n. from Bassus

Zosteragathis dravida (Bhat & Gupta, 1977), comb. n. from Agathis

Zosteragathis elongator (Achterberg & Long, 2010), comb. n. from Therophilus

Zosteragathis festiva (Muesebeck, 1953), comb. n. from Agathis

Zosteragathis festivoides (Sharkey, 1996), comb. n. from Bassus

Zosteragathis fujianicus (Chen & Yang, 2006), comb. n. from Bassus

Zosteragathis gracilis (Bhat & Gupta, 1977), comb. n. from Agathis

Zosteragathis lienhuachihensis (Chou & Sharkey, 1989), comb. n. from Bassus

Zosteragathis lini (Chou & Sharkey, 1989), comb. n. from Bassus

Zosteragathis masoni (Bhat & Gupta, 1977), comb. n. from Agathis

Zosteragathis nigrolineatus (Achterberg & Long, 2010), comb. n. from Therophilus

Zosteragathis nuichuaensis (Achterberg & Long, 2010), comb. n. from Therophilus

Zosteragathis oranae (Watanabe, 1970), (syn. of Z. festiva, syn. by Sharkey, 1996), comb. n. from Agathis

Zosteragathis parasper (Achterberg & Long, 2010), comb. n. from Therophilus

Zosteragathis punctiscutum (Achterberg & Long, 2010), comb. n. from Therophilus

Zosteragathis robusta (Achterberg & Long, 2010), comb. n. from Therophilus

Zosteragathis scutellatus (Achterberg & Long, 2010), comb. n. from Therophilus

Zosteragathis sungkangensis (Chou & Sharkey, 1989), comb. n. from Bassus

Zosteragathis tanycoleosus (Chen & Yang, 2006), comb. n. from Bassus

Zosteragathis samensis Sharkey, sp. n.

Etymology

Named after the type locality Khao Sam Roi Yot National Park.

Diagnosis

Fore coxa yellow. Hind femur black. Second median tergite mostly pale in anterior half and mostly melanic in posterior half. Scutellum sculpture smooth with punctures. Second median tergite dimensions as wide as long or wider.

Figure 18.

Z. samensis: a lateral habitus b fore wing c hind wing d anterior head e lateral head f lateral mesosoma g dorsal mesoscutum h propodeum and median tergites 1-3.

Description

Body length 5.4 mm. Ovipositor length 3.3 mm. Ovipositor 0.6 × body length. Number of flagellomeres 31. Notauli sculpture not significantly wider posteriorly. Scutellum smooth with punctures. Mid tibia with 3 apical and 2 preapical spines. Hind tibia with 8 spines/pegs. Second submarginal cell diameter small, smaller than pedicel length, but larger than pedicel width

Wing hyaline with an infuscate patch posterior to stigma. Second median tergite 0.9 × longer than wide. Second median tergite entirely striate, striae weak anteromedially where they converge medially. Color: head black except gena yellow; mesosoma black; fore and mid coxa yellow; posterior margin of first median tergite yellow; anterior half of second median tergite yellow.

Material examined

Holotype: ♀ (H2418): THAILAND, Prachuab Khiri Khan, Khao Sam Roi Yot NP, Khao Look Glang, 12.107°N, 99.955°E, Malaise trap, 8-15.iii.2009 (H2418), Yai Amnad. Paratypes: All ♀: THAILAND, Prachuab Khiri Khan, Khao Sam Roi Yot NP, foot of Khao Taen, 12.146°N, 99.966°E, 1 m elev., Malaise trap, 3–10.v.2009 (H638, H968), Yai Amnad; Prachuab Khiri Khan, Khao Sam Roi Yot NP, 30 m, N/protection unit4, 12.268°N, 99.944°E, 1 m elev., Malaise trap, 3–10.v.2009 (H973, T4824) 24-31.v.2009 (H490), Yai Amnad; Lampang, Chae Son NP, Youthcamp, 18.83°N, 99.471°E, 455 m elev., Malaise trap, 1–7.iv.2008 (H901) B Kwannui & A. Sukpeng; Mae Hong Son, Namtok Mae Surin NP, Haad Saen, 19.348°N, 97.985°E, Malaise trap, 27.iv–4.v.2008 (H481), Na-maadkam, M; Prachuab Khiri Khan, Khao Sam Roi Yot NP, Saline wetland/Pa Gwad/N, 12.153°N, 99.972°E, Malaise trap, 15–22.iii.2009 (H670), Yai Amnad.

Distribution

Known only from the specimens collected in Thailand. For a distribution map go to: http://bit.ly/1MPrTqu

Acknowledgements

We thank the staff at Queen Sirikit Botanic Garden in Chaing Mai, Thailand for sorting the many hundreds of samples and for the Thai park staff for operating Malaise traps and other collection devices. A special thanks to Chaweewan Hutacharern for managing the Thai end of the TIGER project. Special thanks also to Kees van Achterberg for lending specimens and types from Vietnam. We thank Dr. Nick Stevens and Dr. John Jennings for valuable comments on the manuscript. Funding was provided by NSF grants DEB-0542864 and EF-0337220 and by Hatch projects KY008041 and KY008065 (to MJS). The information reported in this paper (No. 16-08-080) is part of a project of the Kentucky Agricultural Experiment Station and is published with the approval of the Director.

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Appendix I

Specimens used in the phylogenetic analyses, including specimen numbers, GenBank and BOLD accession numbers and rough geographical information.

Taxon name Specimen number Country: Region Type status COI 28S
Aerophilus abdominalis H1313 USA: KY ATRMK294-11 KP943685
Aerophilus malus H1484 USA: WV holotype ATRMK309-11 KP943693
Aerophilus rayfisheri H1212 USA: KY holotype ATRMK278-11 KP943675
Agathacrista depressifera H002 Thailand: Phetchabun KP943596 KC556782
Agathacrista krataei H268 Thailand: Kalasin holotype KP943614 KC556781
Agathacrista sailomi H013 Thailand: Chiang Mai holotype KX431796 KC556780
Agathacrista winloni H502 Thailand: Phetchabun holotype ATRMK218-11 KC771135
Agathigma templei H415 Thailand: Kamphaeng holotype ATRMK211-11 KX431753
Alabagrus maculipes H6020 Mexico: Jalisco ATRMK370-11 KP943698
Asperagathis aspera H274 Thailand: Phetchabun holotype KX431797 KX431706
Asperagathis xesta H095 Thailand: Chaiyaphum holotype KX431798 KX431707
Bassus albifasciatus H014 Thailand: Sakon Nakhon ------ KX431714
Bassus albifasciatus H027 Thailand: Trang KX431800 KX431716
Bassus albifasciatus H032 Thailand: Trang KX431799 KX431715
Bassus albifasciatus H085 Thailand: Trang KX431801 KX431719
Bassus albifasciatus H343 Thailand: Chiang Mai ------ KX431718
Bassus albifasciatus H377 Thailand: Nakhon Si Thammarat ------ KX431717
Bassus alboapicalis H021 Thailand: Trang paratype KX431821 KX431767
Bassus alboapicalis H022 Thailand: Trang paratype KX431819 KX431764
Bassus alboapicalis H081 Thailand: Trang paratype KX431817 KX431762
Bassus alboapicalis H269 Thailand: Trang holotype KX431820 KX431766
Bassus alboapicalis H270 Thailand: Trang paratype KX431818 KX431763
Bassus alboapicalis H307 Thailand: Surat Thani paratype ATRMK195-11 KX431765
Bassus alboapicalis H410 Thailand: Nakhon Si Thammarat paratype ------ KX431761
Bassus albobasalis H003 Thailand: Phetchabun KX431802 KX431721
Bassus albobasalis H092 Thailand: Trang ------ KX431720
Bassus albobasalis H328 Thailand: Phetchabun JQ763436 KX431722
Bassus albocyclus H308 Thailand: Phetchabun paratype ------ KX431724
Bassus albocyclus H349 Thailand: Chiang Mai paratype ------ KX431725
Bassus albocyclus H636 Thailand: Suphan Buri holotype ATRMK230-11 KX431723
Bassus calculator H8008 Sweden: Stockholms län ------ KX431712
Bassus mediatratus H015 Thailand: Chiang Mai holotype KX431816 KX431760
Bassus nopachoni H577 Thailand: Kamphaeng holotype ATRMK223-11 KX431713
Bassus pallidus H055 Thailand: Chanthaburi holotype ------ KX431710
Bassus sp. H376 Thailand: Phetchaburi ATRMK204-11 KX431711
Braunsia smithii H906 Thailand: Chiang Mai ATRMK261-11 HQ667949
Camptothlipsis lingualongis H1887 South Africa: Western Cape paratype ATRMK334-11 JN564494
Camptothlipsis nigra H433 Thailand: Prachuap Khiri Khan ATRMK430-11 HQ667951
Camptothlipsis sheilae H664 Thailand: Kanchanaburi holotype ATRMK235-11 HQ667954
Camptothlipsis sp. H162 Uganda: Homa Bay ------ KX431699
Camptothlipsis sp. H2299 Congo: Pool ------ KX431698
Chimaeragathis chrysoma H710 Thailand: Phetchaburi holotype ATRMK240-11 KX431738
Chimaeragathis eurysoma H045 Thailand: Trang paratype KX431805 KX431736
Chimaeragathis eurysoma H069 Thailand: Trang paratype KX431806 KX431737
Chimaeragathis eurysoma H925 Thailand: Phetchaburi holotype ATRMK265-11 KX431735
Chimaeragathis lohmani H072 Thailand: Trang holotype KX431807 KX431739
Chimaeragathis lohmani H077 Thailand: Trang paratype KX431808 KX431740
Cymagathis krikoma H290 Thailand: Chaiyaphum paratype ATRMK192-11 KX431701
Gyragathis leucosoma H275 Thailand: Nakhon Ratchasima holotype KX431794 KX431700
Leuroagathis paulbakeri H369 Thailand: Prachuap Khiri Khan holotype ------ KX431709
Liragathis baonai H360 Thailand: Nakhon Si Thammarat holotype ATRMK200-11 KX431705
Liragathis damnai H397 Thailand: Chiang Mai paratype ATRMK210-11 KX431704
Liragathis javana H283 Thailand: Trang KX431795 KX431702
Liragathis javana H628 Thailand: Phetchabun ATRMK228-11 KX431703
Neothlipsis parysae H4428 USA: KY paratype ATRMK364-11 KX431696
Neothlipsis sp. H195 Thailand: Surat Thani KP943607 KP943660
Neothlipsis sp. H198 USA: KY KX431793 KX431697
Neothlipsis sp. H7618 Mexico: Yucatan ATRMK403-11 KP943709
Scabagathis emilynadeauae H033 Thailand: Trang holotype KX431792 KX431695
Trochantagathis marshi H067 Thailand: Trang KX431809 KX431742
Trochantagathis marshi H089 Thailand: Trang KX431811 KX431745
Trochantagathis marshi H1851 Thailand: Trang ------ KX431744
Trochantagathis marshi H281 Thailand: Trang KX431810 KX431743
Trochantagathis marshi H765 Thailand: Ubon Ratchathani ATRMK242-11 KX431741
Trochantagathis marshi H799 Thailand: Suphan Buri ------ KX431746
Trochantagathis marshi H965 Thailand: Nakhon Si Thammarat ATRMK266-11 KX431747
Xanthagathis mellisoma H060 Thailand: Trang KX431812 KX431749
Xanthagathis mellisoma H145 Thailand: Phetchabun ------ KX431748
Xanthagathis mellisoma H286 Thailand: Chaiyaphum ATRMK191-11 KX431751
Xanthagathis mellisoma H348 Thailand: Chiang Mai ATRMK199-11 KX431750
Xanthagathis mellisoma H662 Thailand: Phetchaburi ATRMK234-11 KX431752
Zosteragathis contrastus H017 Thailand: Chiang Mai KX431828 KX431783
Zosteragathis contrastus H056 Thailand: Trang KX431834 KX431790
Zosteragathis contrastus H094 Thailand: Chiang Mai KX431833 KX431789
Zosteragathis contrastus H100 Thailand: Chaiyaphum KX431832 KX431787
Zosteragathis contrastus H101 Thailand: Loei KX431827 KX431781
Zosteragathis contrastus H104 Thailand: Loei ------ KX431782
Zosteragathis contrastus H142 Thailand: Nakhon Ratchasima ------ KX431779
Zosteragathis contrastus H143 Thailand: Phetchabun KX431829 KX431784
Zosteragathis contrastus H144 Thailand: Phetchabun KX431830 KX431785
Zosteragathis contrastus H146 Thailand: Phetchabun KX431831 KX431786
Zosteragathis contrastus H149 Thailand: Phetchabun KX431826 KX431780
Zosteragathis contrastus H1855 Thailand: Chaiyaphum ATRMK501-11 ------
Zosteragathis contrastus H603 Thailand: Surat Thani ATRMK226-11 KX431791
Zosteragathis contrastus H677 Thailand: Suphan Buri ------ KX431788
Zosteragathis contrastus H985 Thailand: Kanchanaburi ------ KX431778
Zosteragathis samensis H2418 Thailand: Prachuap Khiri Khan holotype ATRMK475-11 KX431775
Zosteragathis samensis H973 Thailand: Prachuap Khiri Khan paratype ATRMK269-11 KX431774
Zosteragathis sp. H065 Thailand: Trang KX431803 KX431733
Zosteragathis sp. H083 Thailand: Trang KX431804 KX431734
Zosteragathis sp. H091 Thailand: Sakon Nakhon KX443589 KX431726
Zosteragathis sp. H1859 Thailand: Phitsanulok ATRMK329-11 KX431729
Zosteragathis sp. H1860 Thailand: Surat Thani ATRMK330-11 KX431731
Zosteragathis sp. H239 Thailand: Trang ------ KX431732
Zosteragathis sp. H492 Thailand: Phetchaburi ATRMK217-11 KX431728
Zosteragathis sp. H660 Thailand: Mae Hong Son ATRMK233-11 KX431727
Zosteragathis sp. H687 Thailand: Nakhon Si Thammarat ------ KX431730
Zosteragathis sp. H016 Thailand: Chaiyaphum KX431825 KX431776
Zosteragathis sp. H080 Thailand: Chiang Mai KX431814 KX431757
Zosteragathis sp. H121 Thailand: Nong Bua Lam Phu KX431822 KX431771
Zosteragathis sp. H122 Thailand: Nong Bua Lam Phu KX431823 KX431772
Zosteragathis sp. H1625 Thailand: Chaiyaphum ATRMK323-11 KX431754
Zosteragathis sp. H1636 Thailand: Ubon Ratchathani ATRMK325-11 KX431770
Zosteragathis sp. H1858 Thailand: Chiang Mai ATRMK328-11 KX431777
Zosteragathis sp. H236 Thailand: Chiang Mai KX431813 KX431756
Zosteragathis sp. H237 Thailand: Lampang KX431815 KX431758
Zosteragathis sp. H279 Thailand: Ubon Ratchathani KX431824 KX431773
Zosteragathis sp. H473 Thailand: Phetchaburi ATRMK216-11 KX431708
Zosteragathis sp. H598 Thailand: Mae Hong Son ATRMK225-11 KX431768
Zosteragathis sp. H650 Thailand: Phetchabun ATRMK232-11 KX431769
Zosteragathis sp. H689 Thailand: Suphan Buri ATRMK238-11 KX431755
Zosteragathis sp. H989 Thailand: Phetchaburi ATRMK271-11 KX431759