Revision of the Oriental genera of Agathidinae ( Hymenoptera , Braconidae ) with an emphasis on Thailand including interactive keys to genera published in three different formats

Th e genera of Oriental Agathidinae are revised and a fully illustrated dichotomous key is presented. New generic concepts are proposed for Bassus Fabricius, 1804 and Hypsostypos Baltazar, 1963. Bassus is restricted to a clade with an Old World distribution and the remaining members are divided amongst the resurrected genera Camptothlipsis Enderlein, 1920, Lytopylus FÖrster, 1862, and Th erophilus Wesmael, 1837. Th e concept of Hypsostypos is restricted and the new genus Amputostypos Sharkey, gen. n. is proposed to include species formerly included in Hypsostypos that do not have raised antennal bases. Troticus Brullé, 1846 is reported from the Oriental region for the fi rst time. Eighteen genera are recognized for Th ailand and neighboring areas, i.e., Agathis Latreille, 1804, Amputostypos, Aneurobracon Brues, 1930, Bassus, Biroia Szépligeti, 1900, Braunsia Kriechbaumer, 1894, Camptothlipsis, Coccygidium Saussure, 1892, Cremnops Förster, 1862, ZooKeys 21: 19–54 (2009) doi: 10.3897/zookeys.21.271 www.pensoftonline.net/zookeys Copyright M Sharkey et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. Launched to accelerate biodiversity research A peer-reviewed open-access journal


Introduction
Agathidinae is a moderately large subfamily of Braconidae with 1,061 described species worldwide and 238 in the Oriental Region (Yu et al. 2005).Th ough there are an estimated 2,000-3,000 species awaiting description worldwide (Sharkey et al. 2006).Th e subfamily has a worldwide distribution and members are found in most terrestrial habitats.Th ough all known species are koinobiont endoparasitoids of lepidopteran larvae, life history traits vary considerably.Depending on the species, they may be nocturnal or diurnal, gregarious or solitary, attack exposed or concealed hosts, and attack any larval instar.In general they are solitary, attack fi rst-instar Lepidoptera larvae in concealed microhabitats such as leafrolls or stems, and emerge from the last larval instar of the host after it has spun its cocoon.Detailed studies of life history have been conducted for a few species (e.g., Simmonds 1947, Dondale 1954, Odebiyi and Oatman 1972, 1977, Janzen et al. 1998) and a few have been used in classical biological control eff orts.Currently there are about 50 genera recognized (Sharkey 1992).Th e history of higher classifi cation of the Agathidinae was summarized by Sharkey (1992) who also proposed a tribal-level classifi cation based on groundplan coding.Sharkey et al. (2006) conducted phylogenetic analyses based on morphology and the D2-3 regions of 28S rDNA.Perhaps their most noteworthy result was a clear demonstration of the polyphyletic nature of the genus Bassus, however it was beyond the scope of the paper to adjust generic classifi cation.Here we begin to correct this problem by resurrecting two junior synonyms of Bassus and restricted Bassus s.s. to a relatively small Old World clade.
Th e Oriental fauna of Agathidinae were revised by Bhat and Gupta (1977) and they provided a detailed history of taxonomic research for the area.Many of the generic concepts have changed since their publication and here we revise and update the generic concepts and provide a key to genera.Th is is a prelude to a complete revision of all of the species of Agathidinae found in Th ailand funded by a USA, NSF grant to explore the terrestrial insect fauna of Th ailand.As part of the inventory of Th ai insects we have run 3 Malaise traps at 30 diff erent localities throughout Th ailand since 2007, comprising approximately 90 Malaise trap years.

Materials and methods
Material for this study was primarily collected in Malaise traps operated throughout Th ailand since 2007.Raw Malaise trap samples were sent to the Queen Sirikit Botanic Gardens (QSBG) in Chaing Mai, Th ailand, where the Agathidinae and many other taxa were separated.Th ese were then sent to the Sharkey lab in Lexington, Kentucky where they were treated in hexamethyldisilazane, mounted, and labeled.Most species have been sequenced for COI and the D2-3 regions of 28S.Holotypes of all material will be returned to QSBG.For more details on the Th ai inventory visit http://sharkeylab.org/tiger/.
Images for this study were taken with an Automontage© imaging system mounted on a Leica MZ16 stereomicroscope.Th e taxonomic decisions were based primarily on the analyses published in Sharkey et al. (2006) many of which have been confi rmed by more recently obtained unpublished sequence data.
Following the key to genera, below, each genus is treated in alphabetical order.Included in these treatments are discussions of distribution, diversity, phylogeny, biology, and diagnosis.Where warranted, a section on taxonomic decisions is included.Each generic treatment includes a full lateral habitus and an image of a forewing.Th ere are also three published interactive keys available.Th ey are in the following formats: Intkey, Lucid, and MX.
Th e dichotomous key was generated using DELTA software (http://delta-intkey. com).Data, species names, characters and character states were entered into Delta Editor (Dallwitz 1980;Dallwitz et al. 1999).Th e "tokey" fi le was edited to select and weight the characters used for the dichotomous key, and the modifi ed fi le was exported from DELTA to produce the key which was then lightly edited to produce the fi nal version (Dallwitz 1974;Dallwitz 1980;Dallwitz, Paine, and Zurcher 1993).Th e Intkey was produced in a similar manner using the DELTA fi le "toint" and the software Intkey (Dallwitz 1980;Dallwitz, Paine, and Zurcher 1993;Dallwitz, Paine, and Zurcher 1995).All source fi les and images used in this publication are available at http://sharkeylab.org/sharkeylab/ Misc/datasets/DeltaFiles/AgathidinaeTh aiDeltaFiles.zipand in Appendices 1 and 2 of the present paper (doi: 10.3897/zookeys.21.271.app.1.ik and doi: 10.3897/zookeys.21.271. app.2.ik).Th ese fi les are open to the public and future researchers are welcome to download them if they wish to modify, correct, or add newly described taxa for identifi cation.
Online interactive matrix and dichotomous keys were also produced using Lucid (www.lucidcentral.org), and are available on Waspweb at: http://www.waspweb.org/Ichneumonoidea/Braconidae/Keys/index.htm.Users can choose between three diff erent key formats depending on their personal preference: a standard dichotomus key, a Lucid Phoe-nix key or a Lucid matrix key.Lucid Phoenix keys are interactive but still dichotomous and a choice needs to be made at each key couplet to continue.Lucid matrix keys, on the other hand, use a diff erent approach where relevant states from multiple character features can be selected independently until identifi cation is achieved.For more information concerning Lucid keys visit http://www.lucidcentral.org.Files are provided in two formats enabling conversion of the Lucid matrix key to other platforms: 1. Lucid Interchange Format version 3 (LIF3) fi les are XML based fi les that store all the Lucid3 key data, allowing exchange of the key with other key developers, and 2. SDD fi les are XML-based fi les structured using the internationally agreed SDD (Structure of Descriptive Data) Schema.SDD fi les may be used to exchange Lucid keys with other SDD-compliant applications.Lucid fi les are published in Appendix 3 of the present paper (doi: 10.3897/zookeys.21.271.app.3.ik).
A third interactive key to Oriental Agathidinae was generated using MX (Yoder et al. 2006-present) and is available at http://purl.oclc.org/NET/oriental-agathidinae.Th e MX MySQL database fi les are published in Appendix 4 (doi: 10.3897/zookeys.21.271. app.4.ik).Th e MX is optimized for viewing in Firefox.MX is an open source, web-based content management tool and workbench for systematists.Th e images, dynamically generated matrix in Nexus format, and specimen data from the key are available for download.Th e online key has the utility of dynamically updating from database additions.MX has a wide range of functionality beyond multi-entry and dichotomous key generation, information captured from other activities such as managing specimen data, images, phylogenetic characters, and ontologies may be incorporated into key content.Th e key to Oriental Agathidinae is dynamically linked to the Hymenoptera Anatomy Ontology (http://hymao.org), which provides a glossary of terminology for the key.More information regarding MX software is found at http://purl.oclc.org/NET/mx-database.
Th e key data fi les are published under the conditions of the OpenDataCommons license (ODbl) (http://www.opendatacommons.org/licenses/odbl/1.0/) All new species have been registered with Zoobank (Polaszek et al. 2005).Th e concept of data publication and dissemination of interacative keys under the open access model is discussed in a forum paper published in the present issue (Penev et al. 2009).
Distribution: Holarctic, with more diversity in cool temperate regions.No species of Agathis has been collected in Th ailand or in the Oriental Region, but the occurrence of the genus in northern high-altitude areas is likely.Bhat and Gupta (1977) reported 45 species of Agathis for the Oriental region but they used a diff erent generic concept that included Bassus s.s., Th erophilus, and Lytopylus as they are defi ned in the present study.
None of the species treated by Bhat and Gupta (1977) correspond to Agathis s.s., as it is interpreted here.Biology: Species generally attack lepidopterous larvae feeding in fl ower heads.Th ere are numerous host records many of which are likely to be incorrect; host families that are reasonably certain include: Gelechiidae, Coleophoridae, Oecophoridae, Tortricidae, and Prodoxidae.
Diagnosis: Head rostriform or subrostriform (Fig. 17a); tarsal claws not bifi d and with a basal lobe (as in Fig. 2b).Etymology: From the Greek words Amputo and stypos, meaning short and stem respectively.Th ese refer to the close relationship with the genus Hypsostypos, meaning high stem.Amputostypos diff ers from Hypsostypos primarily in lacking high ridges surrounding the antennal bases.

Amputostypos
Taxonomy: Sharkey et al. (2006) included this generic concept under Hypsostypos, mistakenly thinking that the type species of Hypsostypos lacked posterolateral carinae on the frons.
Distribution: Oriental, East Palaearctic, Oceanic, Australian, African (rare), primarily tropical and warm-temperate, but reasonably represented in moderate temperate localities.No specimens are recorded from Th ailand but we have collected 83 specimens representing 10 species.
Diversity: It is diffi cult to estimate the number of species due to recent changes in generic concepts.Sharkey et al. (2006) divided Coccygidium s.l.into Hypsostypos, Zelomorpha, and Coccygidium s.s., however few new combinations were made.Members of Amputostypos are restricted to the Old World and there are about 12 species recorded for the Oriental region.Bhat and Gupta (1977) included 10 species.Amputostylos corresponds to what they referred to as the Sulana species group of Zelomorpha.
Biology: Th ere are no reliable host records available.Th e short ovipositor suggests that they attack exposed hosts.Many species are pale colored with rather large ocelli and presumably nocturnal.
Diagnosis: Members are very similar to Coccygidium and Euagathis.Unlike Coccygidium they have relatively short foretibial spurs (Fig. 4b) and the frons lacks lateral carinae (Fig. 5b).Unlike Euagathis they have one or two carinae ventrally on the hind trochantellus (Fig. 3a).Members diff er from Hypsostypos in lacking the high ridges surrounding the antennal insertions.
Description: Head: Lateral carina of frons lacking (Fig. 5b); interantennal space usually with two weak prominences separated by shallow groove (never high as in Hypsostypos); gena not extended ventroposteriorly into sharp prominence; labial palp with four segments, third segment not reduced, more than half length of apical segment; apical antennomere acute.Mesosoma: Mesoscutum with sculptured notauli; posteroscutellar depression absent; median areola of metanotum surrounded by well defi ned carinae laterally and posteriorly; propodeum areolate carinate; posterolateral corners of propodeum elongate; propleuron mildly convex to fl at; propodeal pseudosternite well developed, separating hind coxal cavities from metasomal foramen.Legs: Foretibial spur not elongate, about ½ length of basitarsus (Fig. 4b); foretibial spur with setae extending to its apex or nearly so (Fig. 4b); foretibia lacking pegs; tarsal claws bifi d (Fig. 2a); midtibia with apical pegs but lacking pegs at midlength; hind femur usually rugose ventrally; hind tibia with 2 apical pegs, posterior peg larger than anterior peg.Wings (Fig. 18b): Rs+Ma vein of forewing incomplete and not tubular throughout; second submarginal cell of forewing triangular and sessile; forewing 3RSb straight to slightly sinuate; hind wing crossvein r absent; hind wing crossvein r-m weakly indicated as a short nebulous or spectral thickening, i.e., as a depressed line that may or may not be pigmented, near the base of Rs; hind wing Cub present as nebulous or spectral vein.Metasoma: All terga smooth, lacking sculpture; median tergite 1 lacking pair of longitudinal carinae; median syntergum 2+3 lacking transverse depression separating terga 2 and 3 or with depression barely indicated; ovipositor, decurved, shorter than half the length of the metasoma when fully extended (Fig. 18a).
Diversity: Five species described world-wide, two recorded for the Oriental region (India, Philippines), and none for Th ailand.No specimens of Aneurobracon have been collected in Th ailand but it is likely that this rare genus occurs in the country.
Biology: Th ere are two host records both on members of the family Gracillariidae.Phylogenetic Information: Sister to Mesocoelus, which is confi ned to the neotropics.
Diagnosis: Th e lack of venation (Fig. 19b), long legs and long setae on the hind tibia (Fig. 19a) are all unique for the Oriental agathidine fauna.
Distribution: Old world tropical, including African, Oriental, and Australian regions.Bhat and Gupta (1977) used the name Isoptronotum for the same concept, and it here considered a junior synonym (following Sharkey et al. 2006).Bhat and Gupta (1977) included 10 species (of the present concept of Biroia) in the Oriental Region.No specimens have been recorded from Th ailand but we have collected one or two species; that is, one polymorphic species or two closely related species.Diversity: 29 species are known of which 12 are recorded from the Oriental region.Biology: Unknown, the long ovipositors suggest concealed hosts.Phylogenetic Information.Sister to Zacremnops, a small genus with a Neotropical distribution (Sharkey et al. 2006).
Taxonomic Information.Most authors treated species under Isoptronotum prior to Sharkey et al. (2006).
Distribution: Old world tropical, including African, Oriental, and Australian regions.Bhat and Gupta (1977) separated the genus Laccagathis, which is here considered a junior synonym (following Sharkey et al. 2006).No specimens are recorded from Th ailand, but we have collected two species.Diversity: 68 species are described world-wide and 26 are known from the Oriental region (Yu et. al. 2005).
Biology: Most host records are on Pyralidae, with one record each for Lasiocampidae and Noctuidae that need confi rmation.
Distribution: Old World, including Palaearctic, African, Oriental, and Australian regions; far more diverse in tropical areas.Related taxa occur in the New World and continued research will determine whether or not these should be considered congeneric.Often included with Bassus s.l. in keys.
Diversity: 17 species have been described, 6 from the Oriental region all of which were included by Bhat and Gupta (1977).No specimens have been recorded from Th ailand but we have collected two species.Th e genus is especially diverse in the Ethiopian region where there are more than 100, mostly undescribed, species.
Biology: Th e three host records are all on Gelechiidae.Phylogenetic Information.In a clade that includes Zacremnops, Plesiocoelus and some taxa presently placed in the polyphyletic Th erophilus (as Bassus s.l. in Sharkey et al. 2006).
Distribution: Oriental, Palaearctic, Oceanic, Australian, African, primarily tropical and warm-temperate, but reasonably represented in moderate temperate localities.No species are recorded from Th ailand but we have collected 18 specimens representing 3 or 4 species in Th ailand.Diversity: It is diffi cult to estimate the number of species due to recent changes in generic concepts.Sharkey et al. (2007) divided Coccygidium s.l.into Hypsostypos, Zelomorpha, and Coccygidium s.s., but did not include a list of new combinations so the generic concepts have not been incorporated into the Taxapad database (Yu et al. 2005).Members of Coccygidium are restricted to the Old World and there are about 10 species recorded for the Oriental region; Bhat and Gupta (1977) included 8 species.Coccygidium corresponds to what they referred to as the Fuliginosa species group of Zelomorpha.
Biology: Th ere are fi ve host records, all on members of the family Noctuidae.Many species are pale colored with rather large ocelli and are nocturnal.Th e short ovipositor suggests that they are attacking exposed hosts.
Phylogenetic Information.Sister to Zelomorpha which is New World and primarily tropical in distribution (Sharkey et al. 2006).
Taxonomic Information.Chou and Sharkey (1989) treated Zelomorpha as a junior synonym; however the monophyly of both Coccygidium and Zelomorpha were confi rmed by Sharkey et al. (2006).
Diagnosis: Th e long, style-like, foretibial spur (Fig. 4a) is unique amongst Agathidinae.Members are otherwise very similar to those of Amputostypos, which are more commonly collected in Malaise traps in the Oriental Region.Diagnosis: Ovipositor longer than half length of metasoma (Fig. 24a); fore and mid tarsal claws cleft (Fig. 2a); notauli impressed (as in Fig. 7b); hind trochantellus lacking ventral carinae (as in Fig. 3b).Biology: Unknown; the long ovipositor suggests that it attacks concealed hosts.Phylogenetic Information.Cremnoptoides is a member of the tribe Cremnoptini but exemplars have not been included in published phylogenetic analyses.Our unpublished analyses of COI and D2-3 28S sequence data place it as sister to Cremnops.
Distribution: Old World, primarily tropical: African, Oriental, and Australian regions, with a few Palaearctic species.No specimens have been recorded from Th ailand but we have collected four species represented by 5 specimens, suggesting that there are considerably more.Diversity: Bhat and Gupta (1977) recorded 23 species from the Oriental region and Bhat (1978) added 2 new Oriental species.
Biology: Most host records are on Noctuidae and the short ovipositors suggest that exposed hosts are attacked.
Phylogenetic Information.Sister to all other Disophrini that were included in the Sharkey et al. (2006) analyses.
Distribution: Holarctic, Oriental, austral region of South America, especially diverse in cold temperate areas.Species described from Chile as Earinus are sister to the Hol- arctic and Oriental clade (Sharkey et al. 2006).Th ere are no records from Th ailand but we have captured one specimen.Diversity: 15 species are described world-wide, and 3 from the Oriental region.Bhat and Gupta (1977) recorded only one species from the Oriental region.Th ere are many undescribed species in Austral South America.
Biology: Most host records are on Noctuidae and Tortricidae.Phylogenetic Information.Sister to all other Earinini (Sharkey et al. 2006;and new unpublished data).
Diagnosis: Th is is the only agathidine genus in the Oriental region with a complete RS+M vein in the fore wing (Fig. 26b).
Biology: Most host records are on Lymantriidae and the short ovipositors suggest that exposed hosts are attacked.
Phylogenetic Information.Rather unplaced within the Disophrini based on Sharkey et al. (2006) although many analyses placed it as sister to Amputostypos (as Hypsostypos in Sharkey et al (2006)) and unpublished COI and 28S sequence data support this placement.
Distribution: Recorded from Sumatra, Peninsular Malaysia, and Yunnan Prov.China.Not recorded from Th ailand but this rare genus undoubted occurs there.
Diversity: 4 described species, 3 species were included in Bhat & Gupta (1977).Biology: No host records but and the short ovipositors suggest that exposed hosts are attacked.
Distribution and Diversity: Th e genus is only represented in the literature by the type species from Sulawesi, though we have seen another specimen representing an undescribed species from Sulawesi.Due to its rarity and proximity to Th ailand it may occur there as well.
Taxonomy: See the Taxonomy section under Amputostypos.Biology: Th e short ovipositor suggests that exposed hosts are attacked.berg 1992, Sharkey 1997), and further showed that a stricter sense of Bassus was a monophyletic group and sister to Braunsia.However they did not examine the type specimen of Bassus which does not happen to belong to the same clade as the specimens included in their analyses.Here we have selected the oldest available name for what was referred to as Bassus s.s. in Sharkey et al. (2006).Lytopylus was fi rst proposed by Förster 1862 but no species were assigned to the genus until Viereck (1914) included L. azygos as the type.Distribution: Cosmopolitan, with more diversity in temperate regions.Only one species of Lytopylus has been collected in Th ailand but the occurrence of more members of the genus is likely.Bhat and Gupta (1977) included members of Lytopylus under Agathis.Th ese include L. aequoreticulatus (Bhat & Gupta, 1977), L. astioles (Nixon, 1950), L. burmensis (Bhat & Gupta, 1977), L. phillipinensis (Bhat & Gupta, 1977) and L. romani (Shestakov, 1940) all new combinations.
Diversity: Highly speciose.Biology: Most commonly attacking species of Tortricidae and Pyralidae, other reliable records include: Elachistidae, Gelechiidae, and Th yrididae.Undoubtedly many other host families will be confi rmed or discovered.
Taxonomy.Based on the results of Sharkey et al. (2006) we have broken the former concept of Bassus into several large monophyletic genera, e.g., Lyptopylus, Camptothlipsis, however not all of the problems with the old concept of Bassus are solved and Th erophilus is here used to contain the polyphyletic assemblage that remains.Clearly there is improvement to be made however it seems better to recognize large monophyletic groups within the former concept of Bassus.If monophyly were the only criterion many agathidine genera would need be lumped into one genus.Diversity: Th is polyphyletic genus is very rich and contains about as many species as all other genera combined.
Biology: Attacking a wide assortment of primarily micro-Lepidoptera, mostly in concealed microhabitats.Undoubtedly many other host families will be confi rmed or discovered.