Revision of the world species of the genus Habroteleia Kieffer (Hymenoptera, Platygastridae, Scelioninae)

Abstract The genus Habroteleia Kieffer is revised. Seven species are recognized, three are redescribed: H. flavipes Kieffer, H. persimilis (Kozlov & Kononova), H. ruficoxa (Kieffer); and four species are described as new: H. mutabilis Chen & Talamas, sp. n., H. salebra Chen & Talamas, sp. n., H. soa Chen & Talamas, sp. n., and H. spinosa Chen & Johnson, sp. n. Four species are treated as junior synonyms of Habroteleia flavipes Kieffer: Chrestoteleia bakeri Kieffer, syn. n., Habroteleia bharatensis Saraswat, syn. n., Habroteleia browni Crawford, syn. n., and Habroteleia kotturensis (Sharma), syn. n. Habroteleia dagavia (Kozlov & Lê), syn. n. is treated as junior synonym of Habroteleia persimilis (Kozlov & Kononova). Baryconus vindhiensis (Sharma), comb. n. is transferred out of Habroteleia Kieffer. Habroteleia impressa (Kieffer) and H. scapularis (Kieffer) remain valid species but their identity and status are unclear.


Introduction
The genus Habroteleia was originally described by Kieffer (1905) based on the type species, Habroteleia flavipes Kieffer, collected on the island of Sumatra, Indonesia. Kieffer (1913) later proposed Chrestoteleia for a single species, Chrestoteleia bakeri Kieffer, collected from the Philippines, which was treated by Baltazar (1961) as a junior synonym of Habroteleia. Nine species have since been described from India, Japan and the Philippines. We here provide the first comprehensive treatment of the genus, including examination of type specimens of all species except H. impressa (Kieffer) and H. scapularis (Kieffer), for which we were unable to locate type material. The previously described species of Habroteleia were recorded from the Oriental region, extending from India to Japan, and we here provide records that expand the distribution of Habroteleia to include Madagascar, Papua New Guinea, and the Fijian archipelago.
The host of Habroteleia is unknown, but we suspect that it parasitizes orthopteran eggs (large and elongate) based on its elongate body and because Orthoptera is suspected to be the plesiomorphic host group for the platygastroids as a whole (Austin et al. 2005).
Morphological terms used in this work were as in the Hymenoptera Anatomy Ontology (Yoder et al. 2010) (Appendix 1). Identifiers (URIs) in the format HAO_ XXXXXXX represent concepts in the HAO and are provided to enable readers to confirm their understanding of the concepts being referenced. To learn more about a given concept, including additional images, notes, references and other metadata, use the identifier as a search term at http://glossary.hymao.org or use the identifier as a web-link.
In the Material Examined section the metadata for the specimens studied are recorded in an abbreviated format, using unique identifiers (numbers prefixed with "OSUC", "CASENT", "FBA", "MNHN_EY") for the individual specimens. The label data for all specimens have been georeferenced and recorded in the Hymenoptera Online database, and details on the data associated with these specimens can be accessed at the following link, hol.osu.edu, and entering the identifier in the form (note the space between the acronym and the number). The electronic version of the paper contains hyperlinks to external resources. Insofar as possible, the external information conforms to standards developed and maintained through the organization Biodiversity Information Standards (Taxonomic Database Working Group). All new species have been prospectively registered with Zoobank (Polaszek et al. 2005, www.zoobank.org), and other taxonomic names, where appropriate, have been retrospectively registered. The external hyperlinks are explicitly cited in the endnotes so that users of the printed version of this article have access to the same resources.
Data associated with the genus Habroteleia can be accessed at http://hol.osu.edu/ index.html?id=488. The generic and species descriptions were generated by a database application, vSysLab (vsyslab.osu.edu), designed to facilitate the production of a taxon by character data matrices, and to integrate those data with the existing taxonomic, media, and specimen-level database. Data may be exported in both text format and as input files for other applications. The text output for descriptions is in the format of "Character: Character state (s)". Polymorphic characters are indicated by semicolonseparated character states.
Images and measurements were produced with multiple systems. Photographs of IEBR specimens were captured with a Canon Rebel 600 camera connected to a Wild M10 microscope with a Fotoprojektiv 2.5×/SLR 10446175 adapter and stacked with the program Zerene Stacker. A scale bar was calibrated for images taken at the maximum magnification of the microscope. The remaining images were produced with Combine ZP and AutoMontage extended-focus software, using a JVC KY-F75U digital camera, Leica Z16 APOA microscope, and 1X objective lens. Images were postprocessed with Abobe Photoshop CS3 Extended. A standard set of images is provided for each species: dorsal habitus, lateral habitus, dorsal and lateral views of the head and mesosoma, and anterior view of head. The individual images are archived in Specimage (specimage.osu.edu), the image database at The Ohio State University.
Images of primary types of H. ruficoxa and H. persimilis were provided by Agnièle Touret-Alby (MNHN) and Konstantin Samartsev (ZIN), respectively. Images of the primary type of Baryconus vindhiensis, Habroteleia bharatensis and Habroteleia kotturensis were made available by  and images of Triteleia dagavia were made available by Talamas and Pham (2017), all are used in this publication with permission.
Diagnosis. Habroteleia can be separated from other scelionines by the combination of the following characters: epomial carina present; malar and facial striae absent; marginal vein many times longer than stigmal vein; postmarginal vein (R1) absent or rudimentary; propodeum with lateral and median projections; T6 in females strongly depressed dorsoventrally to form a flat triangle; male antenna without tyloid (Chen et al. 2013).

Habroteleia flavipes
Comments. This species is well supported by many characters, although the color of mesosoma and metasoma, sculpture of mesoscutal midlobe, and the length of median propodeal projection are variable. The color of mesosoma and metasoma varies from orange to dark brown. The sculpture of the mesoscutal midlobe varies from largely smooth with two rows of discrete coarse punctures to coarsely carinate with two rows of contiguous coarse punctures. The length of the median propodeal projection varies from short to long. These variations are gradual among specimens. Therefore, we consider them as intraspecific rather than interspecific differences.
Comments. The holotype specimen of Habroteleia ruficoxa is in reasonably good condition in that the characters used for diagnosis at the species level are readily accessible. The challenge is that the species was described from a single male and in the course of this revision we did not encounter any additional specimens of H. ruficoxa. The absence of a spine on T8 in the male, the largely smooth surface of the mesos-cutum and mesoscutellum, and the notauli weakly indicated by punctures place the punctate. Length of T6 in female: wider than long. Apex of T6 in female: round. Sculpture of S2: longitudinally striate rugose. Sculpture of T1 in male: sparsely longitudinally striate, smooth in interstices. Male T8 apical spine: present.
Etymology. The Malagasy word "soa" means "beautiful" or "excellent". We apply it to this species because we find it to be both of these. The name is treated as a noun in apposition.
Link to distribution map. Comments. Habroteleia soa is the most geographically disjunct member of the genus, separated from the other species by the Indian Ocean. Despite this separation, it is not morphologically unusual in comparison with the other species, suggesting either that there is a relatively recent division between H. soa and the other species, that the morphology of the genus evolves rather slowly, or that there has been insufficient sampling in the intervening areas (e.g., east Africa, the moist southern part of the Arabian peninsula, India, and all other intervening regions). Description. Body length of female: 3.51-3.52 mm (n=2). Body length of male: 3.37-3.81 mm (n=6). Length of A3 in male: longer than A2. Punctation of frons above antennal scrobe: dense. Sculpture of antennal scrobe: punctate rugose. Central keel: present. Sculpture of ventrolateral frons: punctate rugose. Occipital carina: interrupted medially. Sculpture of posterior vertex: smooth with sparse punctures. Sculpture of gena: sparsely punctate. Sculpture of occiput: smooth.
Color of metasoma: black. T1 horn in female: absent. Sculpture of posterior margin of T1 in female: densely longitudinally striate, punctate rugulose in interstices. Transverse sulcus on T2: present. Sculpture of T2-T5: densely longitudinally striate, punctate rugulose in interstices. Sculpture of T6 in female: rugose. Length of T6 in female: wider than long. Apex of T6 in female: pointed. Sculpture of S2: sparsely longitudinally striate medially, with fine punctures in interstices, irregularly finely punctate laterally. Sculpture of T1 in male: sparsely longitudinally striate, smooth in interstices. Male T8 apical spine: absent. Diagnosis. This species is most similar to H. salebra but can be distinguished by the pointed apex of T6 in females and the spine at the apex of T8 in males, Etymology. The specific epithet means spiny, referring to the pointed apex of T6 in females and should be treated as an adjective.