Revision of the World species of the genus Chromoteleia Ashmead (Hymenoptera, Platygastridae, Scelioninae)

Abstract The genus Chromoteleia Ashmead is revised. Twenty-seven species are recognized, of which six species are redescribed: C.congoana (Risbec), C.connectens Kieffer, C.fuscicornis Kieffer, C.longitarsis Kieffer, C.semicyanea Ashmead, C.tricarinata Kieffer; and twenty-one species are described as new: C.aequalis Chen & Johnson, sp. n., C.alternata Chen & Johnson, sp. n., C.bidens Chen & Masner, sp. n., C.copiosa Chen & Johnson, sp. n., C.cuneus Chen & Johnson, sp. n., C.curta Chen & Johnson, sp. n., C.depilis Chen & Johnson, sp. n., C.dispar Chen & Masner, sp. n., C.feng Chen & Johnson, sp. n., C.fossa Chen & Johnson, sp. n., C.ingens Chen & Masner, sp. n., C.levitas Chen & Johnson, sp. n., C.longa Chen & Johnson, sp. n., C.maura Chen & Masner, sp. n., C.parvitas Chen & Johnson, sp. n., C.pilus Chen & Johnson, sp. n., C.plana Chen & Johnson, sp. n., C.rara Chen & Johnson, sp. n., C.robusta Chen & Johnson, sp. n., C.semilutea Chen & Johnson, sp. n., C.sparsa Chen & Johnson, sp. n.Chromoteleiarufithorax Kieffer remains a valid species, but its identity and status are unclear. All species are known only from the Neotropical region except for Chromoteleiacongoana (Resbec) which only occurs in Africa.


Introduction
The genus Chromoteleia was originally described by Ashmead (1893) based on the colorful type species Chromoteleia semicyanea Ashmead collected in Saint Vincent. Later,  proposed Petalosema for species with expanded metanotum (metascutellum).  synonymized Petalosema with Chromoteleia by pointing out that  was misled by Ashmead's (1893) inaccurate illustration of C. semicyanea showing a narrow and unexpanded metanotum. Six species were recorded from the New World: C. connectens Kieffer, C. fuscicornis Kieffer, C. longitarsis Kieffer, C. rufithorax Kieffer, C. semicyanea Ashmead, and C. tricarinata Kieffer. One species also was described from the Afrotropical region, C. congoana (Risbec), from what is currently Gabon. One fossil species, C. theobaldi Maneval, is reported from Baltic amber (from 40~60 mya) (Maneval 1938), but the status of this species is not clear (Talamas and Buffington 2015) and requires direct examination.
Since its original description in 1893, Chromoteleia has never been comprehensively reviewed. Although there are no host records for Chromoteleia, we suspect that it parasitizes the eggs of Orthoptera based on large size and elongate shape of the parasitoids. A number of scelionine genera of similar habitus, and presumed close relation, are also known to be parasitoids of orthopteran eggs, e.g., Macroteleia Kieffer (Muesebeck, 1977), Triteleia Kieffer (Popovici et al. 2011). The goal of this work is to produce a systematic revision of the world species of Chromoteleia, expand the biogeographic data associated with these species, and to present a clarified generic concept. The contributions of the authors are as follows. H.-y. Chen, E. J. Talamas, A. A. Valerio and N.F. Johnson: character definition, generic concept development, species concept development, imaging, key development, manuscript preparation; L. Masner: character definition, generic concept development, species concept development.
Morphological terminology otherwise generally follows Masner (1980) and Mikó et al. (2007). 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 find out 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 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 de-tails 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 cited explicitly in the endnotes so that users of the printed version of this article have access to the same resources.
Data associated with the genus Chromoteleia can be accessed at http://hol.osu. edu/index.html?id=464. 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 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 made using Combine ZP and AutoMontage extended-focus software, using JVC KY-F75U digital camera, Leica Z16 APOA microscope, and 1X objective lens. Images were post-processed 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.  Kieffer, 1926: 267, 358. Type: Chromoteleia rufithorax Kieffer, by original description (keyed, key to species), designated by Muesebeck and Walkley 1956. Synonymized by ; Muesebeck and Walkley 1956: 336  Inner orbits: diverging ventrally. IOS/EH: IOS distinctly less than EH. Interantennal process: short, often excavate medially. Central keel: present or absent. Antennal foramen: oriented laterally on interantennal process. Facial striae: absent. Malar sulcus: present. Malar striae: absent. Setation of compound eye: absent. Gena: broad, convex, distinctly produced behind eye. Clypeus shape: narrow, slightly convex medially, lateral corners not produced. Anterior (or ventral) margin of clypeus: pointed; straight. Labrum: not visible in anterior view. Number of mandibular teeth: 3. Arrangement of mandibular teeth: transverse. Number of maxillary palpomeres: 4. Shape of maxillary palpomeres: cylindrical. Number of labial palpomeres: 2.

Chromoteleia
Antenna. Number of antennomeres in female: 12. Number of antennomeres in male: 12. Insertion of radicle into A1: parallel to longitudinal axis of A1. Shape of A1: more or less cylindrical, not flattened. Length of A3 of female: distinctly longer than A2. Number of antennomeres with basiconic sensilla in female: 5; 6. Number of antennomeres with basiconic sensilla in female: 5; 6. Arrangement of sensilla on female clava: in longitudinal pairs. Number of antennomeres bearing tyloids in male antenna: 1. Shape of male flagellum: filiform.
Wings. Wing development of female: macropterous. Wing development of male: macropterous. Tubular veins in fore wing: present. Bulla of fore wing R: absent. Length of marginal vein of fore wing: punctiform, R terminating at costal margin. Origin of r-rs in fore wing: basal of point at which R meets costal margin. Basal vein (Rs+M) in fore wing: spectral; nebulous. Development of R vein in hind wing: complete.
Generic diagnosis. The large size and distinctive characters of Chromoteleia (metascutellum large and setose, propodeum without projections, marginal vein of fore wing punctiform) make it a relatively easy genus to identify. The setation of the metascutellum is found in relatively few scelionine genera, typically among the more robust genera, and is a useful diagnostic character. Chromoteleia appears closest to Bracalba Dodd and Romilius Walker, from which it can be separated by the setation of the eyes (absent in Chromoteleia).
Comments. The distribution of Chromoteleia in Africa and South America is a phenomenon of biogeographical interest. Dispersal from South America to Africa has been demonstrated in the parasitoid wasp genus Kapala Cameron (Eucharitidae) (Murray and Heraty 2016) and a similar event could explain the disjunct distribution of Chromoteleia. Alternatively, the dispersal event could have occurred in the opposite direction, followed by radiation in the New World tropics. While there is no direct evidence that the distribution of Chromoteleia represents relictual populations, this is likely the case with other scelionine taxa. For example, Archaeoteleia Masner, which today is known from New Zealand and South America, has been documented from Burmese amber (Talamas et al. 2017), suggesting that the extant fauna of this genus is the remainder of a once widespread distribution.
Chromoteleia is widespread in continental Mesoamerica, Central America, and South America. It is found as far north as the Mexican state of Jalisco, and in the south extends to Itapúa Department in Paraguay and Paraná in southern Brazil. It is noteworthy, though, that it is entirely absent from the Greater Antilles. In the Lesser Antilles, one species, C. semicyanea, apparently is endemic in St. Vincent, and a second, C. aequalis, is known from Dominica (as well as Guyana). This is unusual for scelionines of comparable size and presumed biology: genera such as Scelio, Baryconus, Macroteleia, Triteleia and Opisthacantha are common and richly represented in species throughout the Caribbean.

Basiconic sensilla on A12
Seven of the twenty-seven species of Chromoteleia clearly have two basiconic sensilla on the apical antennomere ( Figure 2), a state that is unknown to us from any other scelionine in which basiconic sensilla are arranged in longitudinal pairs.

Metapleural setation
The metapleuron in Chromoteleia is an important source of characters. In all species there is a line of setigerous foveae along the anterior margin of the metapleuron, corresponding to the dorsal portion of the metapleural sulcus. Directly posterior to this line of setae in the dorsal portion of the sclerite (dorsal metapleural area) there may be patch of setae ( Figure 15, black arrow), and its presence or absence separates numerous species.
In the ventral portion of the sclerite, there is often a line of setae ( Figure 15, white arrow) directly dorsal to the metapleural triangle. These setae can be distinguished from setation of the metapleural triangle because they do not have foveate bases and are located dorsal to the metapleural epicoxal carina, when this carina is present.

Setation of speculum and subalar pit
The presence of setae (Figs 14, 15, blue arrow) on the dorsal speculum and surrounding the subalar pit is found in all species of Chromoteleia and serves as a useful character for generic diagnosis (http://specimage.osu.edu/getImageInfo.html?image_id=89044). However, because this is a newly recognized character, it has yet to be examined thoroughly throughout Scelioninae. This character is known to us from Romilius, Bracalba, and Macroteleia pilosa Muesebeck.

Body color
Ashmead presumably coined the name Chromoteleia in reference to the remarkable metallic blue head and mesosoma in the type species. This turns out to be unique within the genus, but most species are colorful in a different way. Only a small num- ber have the body entirely black or dark brown as is generally typical for other scelionines. Most have more or less extensive parts of the mesosoma colored a rusty red to orange. Masner (1995) and Masner and Hanson (2006) noted that this color pattern is commonly found in a wide range of scelionine genera in species of moderate size (3-10 mm in length) found at lower altitudes (under 2000 m). They attributed the apparent convergent evolution of the color in these taxa to aposematism, a hypothesis that has not yet been critically tested.   (Figs 33,39,97,109,131,137,143,149,155,173,209,237,238 Metasoma variably patterned in alternating orange-yellow and dark brown ( Figure 41); mesoscutellum smooth medially, densely punctate laterally (Fig-ure39)  Occiput rugose ( Figure 97); area directly dorsal to the metapleural triangle with setae ( Figure 95); hind coxa densely punctate ( Figure 95); T6 at least 1.5× longer than wide (Figure 19) .   (Figs 22,57,75,87,93,105,117,169,181,187,205,217,223       Metascutellum with deeply incised apex, forming two spines laterally (     Diagnosis. This species is easily recognized by its entirely black body both in female and male, densely punctate mesoscutum, and abbreviated median mesoscutal carina.
Etymology. The epithet is inspired by the Latin word for equal, in reference to the black body color shared by males and females, and is intended to be treated as an adjective.
Link to distribution map. Transverse sulcus on T2: present. Sculpture of T2: densely longitudinally striate, punctate rugulose in interstices. Sculpture of T6 in female: longitudinally punctate rugose. Length of T6 in female: at least 1.5× longer than wide. Shape of T6 in female in lateral view: flat. Apical spine on female T6: absent. Sculpture of T6 in male: densely punctate. Sculpture of T7 in male: densely punctate. Posterior margin of T7 in male: straight. Sculpture of medial S2: densely longitudinally striate with fine punctures in interstices.
Diagnosis. This species can be easily distinguished from other Chromoteleia species by the following combination of characters: female metasoma variably patterned in alternating orange yellow and dark brown, central keel present only in ventral portion of frons, and notaulus smooth.
Etymology. The epithet is inspired by the Latin word for alternate, in reference to the variably patterned in alternating orange yellow and yellow on metasoma, and is intended to be treated as a noun.

Chromoteleia bidens
Etymology. The epithet is inspired by the Latin word for two-toothed, in reference to the bispinose metascutellum, and is intended to be treated as a noun.
Comments. This species exhibits variation in color and microsculpture. The development of microsculpture on vertex could be absent or present; mesoscutum smooth or carinate medially; mesoscutellum, mesopleuron, and metapleuron ranging from orange to dark brown or black. Diagnosis. This species is similar to C. sparsa, but it can be distinguished by its medially interrupted occipital carina, transversely striate netrion, and the hind basitarsus that is distinctly longer than the remaining segments combined.
Etymology. The epithet is inspired by the Latin word for abundant, in reference to the densely punctate rugose ventral mesepisternum, and is intended to be treated as an adjective.
Diagnosis. This species can be distinguished from other Chromoteleia by the following combination of characters: frons with central keel developed only in ventral portion of frons, dorsal metapleural area without setae, postmarginal vein distinctly shorter than stigmal vein, apex of T6 acute in dorsal view, male T7 with posterior margin deeply emarginate medially between rounded projections.
Etymology. The epithet is inspired by the Latin word for wedge, in reference to the shape of T6 in female, and is intended to be treated as a noun.
Link to distribution map. Diagnosis. This species is similar to C. parvitas, but it can be distinguished by the combination of the following characters: A5 in female distinctly longer than wide, postmarginal vein approximately as long as stigmal vein, female T1 horn present.
Etymology. The epithet is inspired by the Latin word for shortened, in reference to the short median mesoscutal carina, and is intended to be treated as an adjective.
Link to distribution map. Diagnosis. This species is similar to C. pilus, but it can be distinguished by the combination of the following characters: dorsal metapleural area without setation, A6 in female distinctly longer than wide.
Etymology. The epithet is inspired by the Latin word for hairless, in reference to the hairless dorsal metapleural area, and is intended to be treated as an adjective.
Etymology. The epithet is inspired by the Latin word for different, in reference to the different colors of metasoma between female and male, and is intended to be treated as an adjective.
Etymology. The epithet is inspired by the Chinese word 锋 (fēng) for the name of the sharp point of a spear, in reference to the pointed female T6, and is intended to be treated as a noun.
Diagnosis. This species is similar to C. feng with which it shares the characters of smooth notaulus and female A12 with one basiconic sensillum, but it can be distinguished by the combination of the following characters: female T6 without apical spine, occipital carina interrupted medially.
Etymology. The epithet is inspired by the Latin word for ditch, in reference to the smooth notaulus, and is intended to be treated as a noun.
Length of postmarginal vein: distinctly shorter than stigmal vein. Color of metasoma in female: black. Color of metasoma in male: black. Horn on T1 in female: present. Striae of posterior margin of T1 in female: dense. Striae of T1 in male: sparse. Transverse sulcus on T2: absent. Sculpture of T2: densely longitudinally striate, punctate rugulose in interstices. Sculpture of T6 in female: densely longitudinally striate, with fine punctures in interstices. Length of T6 in female: at least 1.5× longer than wide. Shape of T6 in female in lateral view: sinuate. Apical spine on female T6: absent. Sculpture of T6 in male: densely longitudinally striate with fine punctures in interstices. Sculpture of T7 in male: smooth anteriorly, rugulose posteriorly. Posterior margin of T7 in male: emarginate medially between rounded projections. Sculpture of medial S2: densely longitudinally striate with fine punctures in interstices.
Diagnosis. This species can be distinguished from other Chromoteleia by the following combination of characters: female A7 with one basiconic sensillum, dorsal metapleural area with setae, basitarsus distinctly longer than remaining segments combined, postmarginal vein distinctly shorter than stigmal vein, male T7 with posterior margin deeply emarginate medially between rounded projections.

Chromoteleia ingens
Length of postmarginal vein: distinctly longer than stigmal vein. Color of metasoma in female: black. Color of metasoma in male: black. Horn on T1 in female: present. Striae of posterior margin of T1 in female: dense. Striae of T1 in male: sparse. Transverse sulcus on T2: present. Sculpture of T2: longitudinally punctate rugose. Sculpture of T6 in female: densely longitudinally striate, with fine punctures in interstices. Length of T6 in female: at least 1.5× longer than wide. Shape of T6 in female in lateral view: flat. Apical spine on female T6: absent. Sculpture of T6 in male: densely longitudinally striate with fine punctures in interstices. Sculpture of T7 in male: smooth to coriaceous. Posterior margin of T7 in male: emarginate medially between rounded projections. Sculpture of medial S2: densely longitudinally striate with fine punctures in interstices.
Diagnosis. This species is similar to C. longitarsis, but it can be distinguished by the combination of the following characters: female A12 has two basiconic sensilla, and occipital carina complete.
Etymology. The epithet is inspired by the Latin word for huge, in reference to the large body size of this species, and is intended to be treated as an adjective.
Diagnosis. This species is similar to C. tricarinata, but it can be distinguished by the combination of the following characters: female A6 distinctly longer than wide, female A12 with one basiconic sensillum, median mesoscutal carina present anteriorly.
Etymology. The epithet is inspired by the Latin word for dark, in reference to the black body and legs, and is intended to be treated as an adjective.
Link to distribution map. Diagnosis. This species is similar to C. curta, but it can be distinguished by the combination of the following characters: A5 in female as long as wide, postmarginal vein distinctly shorter than stigmal vein, female T1 horn absent.
Etymology. The epithet is inspired by the Latin word for small, in reference to the small body size of this species, and is intended to be treated as a noun.
Length of postmarginal vein: approximately as long as stigmal vein. Color of metasoma in female: black. Horn on T1 in female: present. Striae of posterior margin of T1 in female: dense. Transverse sulcus on T2: present. Sculpture of T2: densely longitudinally striate, punctate rugulose in interstices, with a narrow smooth strip medially. Sculpture of T6 in female: densely longitudinally striate, with fine punctures in interstices. Length of T6 in female: at least 1.5× longer than wide. Shape of T6 in female in lateral view: flat. Apical spine on female T6: absent. Sculpture of medial S2: punctate rugose, with a narrow smooth strip medially.
Diagnosis. This species can be easily distinguished by the following characters: body extremely elongate, central keel present only in ventral portion of frons, mesoscutellum strongly transverse, and T2-T6 with with a narrow smooth strip medially.
Etymology. The epithet is inspired by the Latin word for flat or plain, in reference to the flat and smooth mesoscutellum, and is intended to be treated as an adjective.
Link to distribution map.   Comments. This species is extremely variable in body length, which may be an indication that it attacks multiple species with different size eggs, or that the eggs of its host(s) vary widely in size.
Etymology. The epithet is inspired by the Latin word for strong or robust, in reference to the robust appearance of this species, and is intended to be treated as an adjective.
Link to distribution map. Color of mesosoma: variably orange to nearly black. Sculpture of epicoxal lobe posterior of propleural epicoxal sulcus: densely punctate. Sculpture of lateral pronotal area above pronotal cervical sulcus: smooth dorsally, rugose ventrally. Sculpture of netrion: rugose. Microsculpture of mesoscutum: absent. Macrosculpture of mesoscutal midlobe: punctate rugose anteriorly, sparsely punctate posteriorly. Macrosculpture of lateral lobe of mesoscutum: sparsely punctate. Sculpture of notaulus: foveate. Notaular foveae: interconnected. Median mesoscutal carina: present anteriorly, not extending to posterior margin of mesoscutum. Mesoscutellum in lateral view: flat. Comments. This species is extremely variable in mesosoma color and can be divided roughly into dark and light forms. long. Shape of T6 in female in lateral view: flat. Apical spine on female T6: absent. Sculpture of T6 in male: densely punctate. Sculpture of T7 in male: granulate. Posterior margin of T7 in male: straight. Sculpture of medial S2: densely longitudinally striate with fine punctures in interstices.
Diagnosis. This species can be distinguished from other Chromoteleia by the following combination of characters: female A7 without basiconic sensillum, mesoscutum without median mesoscutal carina, dorsal metapleural area with setae, ventral metapleural area obliquely striate, male T7 with posterior margin straight.
Florida Department of Agriculture and Consumer Services -Division of Plant Industry for their support on this contribution. This material is based upon work supported in part by the National Science Foundation of USA under grant No. DEB-0614764 to N.F. Johnson and A.D. Austin.