Revision of Zosteragathis Sharkey of Thailand (Hymenoptera, Braconidae, Agathidinae, Agathidini)

Based on cladistic analyses recently conducted by Sharkey and Chapman, the genus Zosteragathis Sharkey is revised. Twenty-two species are reported from Thailand, three previously described species, Z. samensis, Z. contrasta and Z. nuichuaensis, and 19 new species, i.e., Z. chaiyaphumensis, Z. eukos, Z. hinensis, Z. hongensis, Z. inthanonensis, Z. krachanensis, Z. lampangensis, Z. lampooensis, Z. luangensis, Z. ngamensis, Z. perknos, Z. petchaburiensis, Z. phahompokensis, Z. phuphanensis, Z. sakaeratensis, Z. sakonensis, Z. samensis, Z. surinensis, Z. taemensis, Z. tonensis. Members of Zosteragathis are known from the Australian, Ethiopian, Oceania, Oriental, and eastern Palaearctic regions.


Morphological terms:
The length of the first metasomal median tergite is measured from the apex of the tendon emanating from the propodeum to the posterior border of the tergite.Metasomal median tergites are abbreviated as follows, T1 = metasomal median tergite 1, T2 = metasomal median tergite 2. T2-3 = metasomal median syntergite 2 + 3. Other terms are from Sharkey and Wharton (1997).Morphological terms used in this revision can be found in the Hymenoptera Anatomy Ontology (HAO; Yoder et al. 2010).To find definitions for any structure search for the term at http://glossary.hymao.org.

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

Species description format:
Descriptions are of the holotype; variation is given in parentheses.Color is not extensively described because the images serve this purpose; however color characters that are variable or of diagnostic significance are detailed.All species are treated with a diagnosis and distributional data.They are illustrated with color photos using a JVC digital camera mounted on a Leica MZ16 microscope and Automontag-eR stacking software.
Species delimitation: We used evidence from molecular data (COI and 28S) and morphology to arrive at species concepts.Details are given in Sharkey and Stoelb (2013).Table 1 gives details on the COI distances within species of Zosteragathis as well as distances to the nearest species.
Specimen collection: As part of the inventory of Thai insects, three Malaise traps at each of 30 different localities throughout Thailand were operated from 2007-2010, com-prising approximately 90 trap-years.The specimens dealt with here are primarily from these traps.Two numbers, beginning with the letters "H" and "T" are listed for each specimen collected by our collaborators in Thailand.The H-number is the specimen number.Specimen information is stored in the Symbiota database (Gries et al. 2014) under the Hymenoptera Institute Collection (HIC).(Symbiota Collections of Arthropods Network; http://symbiota4.acis.ufl.edu/scan/portal/).To search for a specimen in the database, do the following: Under the Search tab (upper left of the screen), select Search Collections.Deselect all collections, and scroll down the Southeast section, put a check in the box next to Hymenoptera Institute Collection, scroll back up and hit the Search > button (right side of screen).H-numbered specimens are stored with a 4-letter prefix (HICH) followed by a 6-digit number.Therefore, H660, as it appears in this publication, is stored as HICH000660.To search for this specimen, scroll down to the Specimen Criteria section, type this number in the box next to Catalog Number and hit the search button.This displays a page with a summary of the specimen information.Clicking on Full Record Details opens a new window with the full specimen record, including all available images.The T-number is the number associated with a single Malaise trap for a single trapping period (usually one week).A complete list of these numbers and associated collection events is available from the authors upon request.
Links to species maps in this paper were generated from the Symbiota database.These are not static maps and as georeferenced specimens are added to the database, the maps will update in real time such that a reader  following a map link will see a map of all of the specimens of the taxon at hand in the database, including those that may have been added after this paper is published.
Clicking on the dots on these maps links to the specimen record in the database.

Phylogenetic methods:
The data set and analyses presented herein is from Sharkey and Chapman (2017a).In that paper, the Zosteragathis terminals were largely unnamed, leaving species descriptions for this paper.In short, we conducted maximum likelihood (ML) phylogenetic analyses on a concatenated COI + 28S dataset (1,313 total characters) using Garli (v. 2.01;Zwickl 2006).The 28S multiple alignment was assembled using the default settings on the MAFFT server (http://mafft.cbrc.jp/alignment/server/;v. 7; Katoh et al. 2006), and employing the Q-INS-i strategy which takes secondary RNA structure into account.All 28S bases were included in subsequent phylogenetic analyses.The data were partitioned by gene region and codon position (COI: 3 partitions; 28S: un-partitioned, total of 4 partitions).The most complex model available (GTR+I+G; Rodriguez et al. 1990) was applied to each partition as per recommendations of Huelsenbeck and Rannala (2004).A 20-replicate ML search for the tree of highest log-likelihood and a 500-replicate ML bootstrap analysis (Felsenstein 1985) were conducted using the default settings.Outgroup genera (Alabagrus, Braunsia, Aerophilus, Camptothlipsis and Neothlipsis) were selected based on two criteria: (1) they comprise genera that are members of three clades that are successively sister to the ingroup clade in the phylogeny in Sharkey and Chapman (2017b), and (2) full length sequences of COI and 28S were available for most of these OTUs.The data set analyzed herein is available from the authors upon request.All of the specimens for which we have DNA sequences are listed in Appendix I, including their accession numbers.
Phylogenetic considerations: Sharkey and Chapman (2017b) revised the agathidine tribes and showed that Zosteragathis falls within the Agathidini.They could not demonstrate the monophyly of Zosteragathis, but confounded with the choice of erecting four genera for morphologically uniform clades, or one genus for the lot, they opted for the latter.The cladogram in Figure 1 shows that none of basal clades that separate the four Zosteragathis clades have support, therefore it is entirely possible that the genus is monophyletic.

Results and discussion
Below is a discussion of the taxonomy of Zosteragathis, a key to the Thai species of this genus, and descriptions of each species.

Taxonomy
Zosteragathis Sharkey, 2017 Type species.Zosteragathis samensis Sharkey, 2017 Diagnosis.Most species can be recognized as members of this genus by the combination of finely microsculptured striae on T2 that end abruptly just anterior to the posterior margin of the tergite; this in combination with a pale colored band in the anterior half of T2, or T2 entirely black.Some species have reduced striae on T2 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; T1 without prominent lateral carinae or medial carina; fore tarsus without spines or pegs.
Etymology.From the Greek zoster, meaning "belt or girdle" and agathis the type genus of the subfamily.Zoster is a reference to the diagnostic pale colored transverse band that is usually present on T2.The gender is feminine.
Color. head black except gena yellow; mesosoma black; fore and mid coxa black; T1 entirely black; anterior margin of T2 yellow.
Etymology.Named after the type locality Chaiyaphum Provence.lomeres 34 (32-39).Sculpture of notauli slightly increasing in width posteriorly but not extending onto lateral lobes of mesoscutum.Scutellum smooth with punctures.Mid tibia with 2 apical and 3 preapical spines.Hind tibia with 7 spines/pegs.Second submarginal cell diameter small, not much larger than width of pedicel.Wing hyaline, without distinct infuscate areas.T2 1× longer than wide.T2 mostly smooth with hints of longitudinal striae especially medially.Color: head black except lower gena yellow; pro and mesothorax orange except ventral mesopleuron melanic; metathorax and propodeum melanic; fore and mid coxae mostly yellow; apex of T1 and lateral and anterior margins of T2 yellow.
For a map of examined material, see: https://bit.ly/2q4eDp2 Distribution.Vietnam and Thailand.
Diagnosis.T2 almost entirely melanic and longer than wide; fore wing lacking infuscate patch posterior to stigma; similar to Z. annuliferus (Achterberg and Long 2010) but dimensions of T1 and T2 differ.
Description.Body length 4.9 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 indistinct carinae that approach the median ocellus.Antenna with 32 flagellomeres.Third labial (penultimate) palpomere minute, barely visible and much smaller than apical palpomere.Scutellar groove with 3 longitudinal ridges.Fore tibia lacking thickened spines; mid tibia with 3 pegs; hind tibia with 10 pegs.
Specimens examined.Diagnosis.Pronotum melanic dorsomedially, concolorous with lateral surfaces of pronotum.Scutellum sculpture smooth with punctures.Description.Body length 4.7 mm.Ovipositor length 3.7 mm.Ovipositor 0.8× body length.Number of flagellomeres 34.Notauli sculpture not significantly wider posteriorly.Scutellum smooth with punctures.Mid tibia with 2 apical and 3 preapical spines.Hind tibia with 6 spines/ pegs.Second submarginal cell diameter small, slightly larger than pedicel width (varying to minute and about the same size as width of pedicel).Wing hyaline with an infuscate patch posterior to stigma.T2 1.1× longer than wide.T2 entirely striate, striae relative straight throughout.Color: head black; mesosoma black; fore and mid coxa yellow; posterior border of T1 yellow; T2 yellow anteriorly and anterolaterally (varying to yellow in entire anterior half).
Description.Body length 6.7 mm.Ovipositor length 6.4 mm.Ovipositor 1.0× body length.Number of flagellomeres broken after the 33rd flagellomere.Sculpture of notauli increasing in width posteriorly where it extends over onto lateral lobes of mesoscutum.Scutellum rugose.Mid tibia with 1 apical and 2 preapical spines.Hind tibia with 7 spines/pegs.Second submarginal cell diameter large, about as large as pedicel length.Wing hyaline with weak infuscation posterior to stigma.T2 2× longer than wide.T2 entirely and evenly striate, striae diverging anteriorly, otherwise approximately straight.Color: head and mesosoma melanic, except lower gena and clypeus yellow; fore and mid coxae yellow; T1 mostly pale yellow with a pair of large melanic spots; T2 entirely pale yellow.
Etymology.Named after the type locality Kaeng Krachan National Park.
Color.Head and mesosoma melanic, except tegula and lowermost extremity of gena yellow; fore coxa yel-     Description.Body length 4.1 mm.Ovipositor length 2.0 mm.Ovipositor 0.5× body length.Number of flagellomeres 34.Sculpture of notauli increasing in width posteriorly where it extends over onto lateral lobes of mesoscutum.Scutellum rugose.Mid tibia with no apical and 2 preapical spines.Hind tibia with 5 spines/pegs.Second submarginal cell diameter large, about as high as pedicel.Wing hyaline, apical half of wing weakly infuscate, as is area posterior to stigma (infuscate areas may not be evident unless lighting is optimal).T2 1.2× longer than wide.T2 entirely striate, striae relative straight throughout.Color: head, prothorax and mesothorax yellow, metathorax and propodeum black; fore and mid coxae yellow; basal 1/3 and apical margin of T1 yellow; anterior 1/2 of T2 yellow.
Etymology.Named after the type locality Pa Hin Ngam National Park.
Description.Body length 3.2 mm.Ovipositor length 2.7 mm. Ovipositor 0.8× body length.Number of flagellomeres (26-28 in Thai specimens) incomplete in holotype.Sculpture of notauli increasing in width posteriorly where it extends over onto lateral lobes of mesoscutum.Scutellum rugose.Mid tibia with 2 apical and 3 preapical spines.Hind tibia with 6 spines/pegs.Second submarginal cell diameter large, almost as large as pedicel length.Wing hyaline, without distinct infuscate areas.T2 0.9× longer than wide.T2 mostly smooth with striae indicated especially posteromedially (to entirely striate, with striae weaker medially), striae relatively straight.Color: head black except gena yellow ventrally; fore and mid coxae mostly melanic; apex of T1 and anterior half of T2 yellow.
Etymology.Perknos is Greek for dusky; here it is a reference to the dark color of the base of the hind tibia.
Etymology.Named after the type locality Petchaburi Provence.
Description.Body length 4.1 mm.Ovipositor length 3.2 mm.Ovipositor 0.8× body length.Number of flagellomeres 31.Notauli sculpture not significantly wider posteriorly.Scutellum smooth with punctures.Mid tibia with 2 apical and 4 preapical spines.Hind tibia with 7 spines/ pegs.Second submarginal cell diameter large, about same size as pedicel length (to 1/3 length of pedicel but always wider than pedicel).Wing hyaline, without distinct infuscate areas.T2 0.9× longer than wide.T2 mostly smooth with short longitudinal carinae restricted to area near median transverse depression.Color: head black except ventral gena and ventral clypeus yellow; mesosoma black; fore and mid coxa yellow; T1 yellow with large black patch posteromedially; T2 yellow in anterior half.
Color.Head black except ventral extremity of gena yellow; mesosoma black; fore and mid coxa yellow; posterior margin of T1 yellow; T2 yellow anteriorly and anterolaterally.
Etymology.Named after the type locality Phu Phan National Park.
Etymology.Named after the type locality Sakaerat Experimental Station.
Description.Body length 3.6 mm.Ovipositor length 3.0 mm.Ovipositor 0.8× body length.Number of flagellomeres 29.Notauli sculpture not significantly wider posteriorly.Scutellum smooth with punctures.Mid tibia with 2 apical and 3 preapical spines.Hind tibia with 6 spines/ pegs.Second submarginal cell diameter minute, about equal to width of vein.Wing hyaline with an infuscate patch posterior to stigma.T2 weakly striate, striae almost absent anteromedially, weakly converging in posterior half.Color: head black except clypeus and lower gena yellow; mesosoma black; fore and mid coxae yellow; T1 and T2 predominantly pale yellow, T1 with a large melanic spot in basal half, T2 with melanic infusion posteromedially.
Etymology.Named after the type locality Sakon Nakhon Provence.
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.T2 0.9× longer than wide.T2 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 T1 yellow; anterior half of T2 yellow.
Etymology.Named after the type locality Khao Sam Roi Yot National Park.
Material examined.Holotype: Female: Thailand, Prachuab Khiri Khan, Khao Sam Roi Yot NP, Khao Look       Scutellum smooth with punctures.Mid tibia with 3 apical and 3 preapical spines.Hind tibia with 7 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.T2 1.1× longer than wide.T2 entirely striate, striae relatively straight.Color: head black except ventral gena yellow; mesosoma black; fore and hind coxa black; posterior margin of T1 yellow; T2 mostly yellow in basal half with melanic tones posteromedially.
Etymology.Named after the type locality Tat Tone National Park.

Figure 1 .
Figure 1.Tree of highest log-likelihood from 20 ML search reps of a combined COI+28S data set with bootstrap values ≥50% (500 search replicates) plotted at the nodes.Tree modified from Sharkey and Chapman (2017a).

Table 1 .
COI divergence data COI data for each species.If a species has more than one terminal, both intra-and interspecific uncorrected p-distances are reported.Interspecific p-distances are to the nearest OTU in the COI tree (not shown).Values are averaged if two or more comparisons are made.

Zosteragathis tonensis Sharkey, sp. n.
pegs.Second submarginal cell diameter minute, about equal to width of vein.Wing hyaline with an infuscate patch posterior to stigma.T2 0.9× longer than wide.T2 entirely striate, semicircular pattern anteromedially, striae converging posterior to this pattern.Color: head black except ventral gena and ventral clypeus yellow; mesosoma black except scutellar groove pale; fore and mid coxae predominantly melanic; T1 black except posterior margin yellow; T2 yellow in anterior half.