The Phanuromyia galeata species group (Hymenoptera, Platygastridae, Telenominae): shining a lantern into an unexplored corner of Neotropical diversity

Abstract The Phanuromyia galeata species group is delineated and its species richness explored for the first time (Hymenoptera: Platygastridae, Telenominae). Fifteen species are described, all of which are new: Phanuromyia comata Nesheim & Masner, sp. n. (Brazil), P. constellata Nesheim, sp. n. (Paraguay), P. corys Nesheim & Masner, sp. n. (Brazil), P. cranos Nesheim & Masner, sp. n. (Bolivia, Costa Rica, Ecuador, French Guiana), P. cudo Nesheim & Masner, sp. n. (Belize, Bolivia, Brazil, Colombia, Costa Rica, Ecuador, French Guiana, Panama, Peru, Trinidad and Tobago, Venezuela), P. dissidens Nesheim & Masner, sp. n. (Bolivia, Brazil, French Guiana), P. galeata Nesheim & Masner, sp. n. (Belize, Brazil, Colombia, Costa Rica, Ecuador, El Salvador, French Guiana, Mexico, Peru), P. galerita Nesheim & Masner, sp. n. (Brazil, Ecuador, French Guiana), P. hjalmr Nesheim, sp. n. (Bolivia, Costa Rica, Ecuador, Paraguay, Venezuela), P. krossotos Nesheim, sp. n. (Ecuador), P. odo Nesheim, sp. n. (Belize, Bolivia, Brazil, Colombia, Costa Rica, Ecuador, El Salvador, French Guiana, Guatemala, Mexico, Panama, Peru, Trinidad and Tobago, Venezuela), P. pauper Nesheim, sp. n. (Ecuador, Peru), P. princeps Nesheim, sp. n. (Brazil, Ecuador, French Guiana), P. tonsura Nesheim, sp. n. (Brazil, Colombia, Ecuador, Paraguay, Peru), P. tubulifer Nesheim & Masner, sp. n. (Brazil, Guyana).


Introduction
The subfamily Telenominae (Hymenoptera: Platygastroidea, Platygastridae) is a large group of egg-parasitoid wasps, comprising 905 known species found throughout the world. Traditionally it has been thought to be composed of two major genera, Telenomus Haliday and Trissolcus Ashmead, and a number of small, morphologically distinctive satellite genera. This view of telenomine diversity was largely based on over 180 years of work on the Holarctic fauna, beginning with Haliday (1833). This scheme, however, has struggled to cope with the diversity of species from the world's tropics.
The genus Phanuromyia Dodd was originally described in 1914 on the basis of a single species from southern coastal Queensland, distinguishing it on the basis of the presence of an extruded ovipositor. Dodd later (1916) added a second species from New South Wales. In the years that followed the concept of the genus largely fell into obscurity, being cited only five times in the taxonomic literature for the rest of the century, largely because Dodd's laconic description did little to distinguish the taxon from the many other species in the subfamily and because the original type material in Australia was not studied. Johnson and Musetti (2003) sought to better define the genus, using new characters and a perspective informed by decades of new collections from around the world. Mineo (2006) rejected the recognition of Phanuromyia, basing this on the a priori assertion that the characters used were only appropriate for distinguishing species groups and not genera. Taekul et al. (2014), however, confirmed that Phanuromyia is distinct from Telenomus and, further, expanded the concept to embrace species before placed in the crassiclava group of Telenomus (following Johnson 1984). The data and analyses (Taekul et al. 2014) suggest that Phanuromyia is the sister group of Telenomus+Trissolcus (along with several satellite genera), and the limited host data that are available all indicate that Phanuromyia are egg parasitoids of lanternflies and planthoppers in the families Fulgoridae and Flatidae (Hemiptera: Auchenorrhyncha).
Information management. Holotypes are unambiguously identifiable by means of the unique identifier or the red holotype label. The numbers prefixed with "OSUC " are unique identifiers for the individual specimens. These unique identifiers are associated with their specimens' data in The Ohio State University's Hymenoptera Online database, which can be accessed at http://hol.osu.edu. Searching this database using a specimen's unique identifier will produce all data associated with the specimen. All new species have been prospectively registered with Zoobank as well as the Hymenoptera Name Server (http:// hns.osu.edu).
Tools. Images were created using AutoMontage and Combine ZP extended focus software. All images are archived within The Ohio State University's image database (http://specimage.osu.edu). Species descriptions were generated using a database application, vSysLab (http://vsyslab.osu.edu). This application facilitates the construction of taxon character data matrices, the integration of matrices with our existing taxonomic database, and the exportation of data in a variety of file types which can be used in other programs.
Species concept. We define species as populations with the potential to interbreed (Mayr 1942). Interbreeding populations will develop a gradient of character states within each morphological character, while distinct separations between character states will exist in non-interbreeding populations (Wild 2004); therefore, species delimitations are made by identifying discrete character states within characters that are present across multiple specimens. Diagnosis. The three genera Phanuromyia, Telenomus and Trissolcus Ashmead cannot be distinguished on the basis of any single, easily recognized morphological character. Rather, they are recognized by the preponderance of evidence from several characters: presence or absence of sculpture on the medial portion of the frons, length of setation between the ommatidia of the compound eyes, shape of the head, number of clavomeres in the female antenna, presence or absence of notauli on the mesoscutum, presence or absence of sculpture on the disk of the mesoscutellum, form of the sternaulus, shape of the first and second metasomal tergites, and sculpture of the second metasomal tergite. To distinguish Phanuromyia, focus should first be placed on the sternaulus. In the large majority of species this is expressed as a line of pits, beginning anteriorly on the mesepisternum near the dorsal apex of the acetabular carina and extending dorsally and posteriorly toward the mesopleural pit. In Telenomus and Trissolcus the sternaulus may have a single irregularly shaped pit, and its course is otherwise represented by fold or crease in the cuticle. Small individuals of Phanuromyia, however, also may have merely a poorly defined line of impression. Supplemental characters to distinguish Phanuromyia are eye setation very short or seemingly absent; frontal depression weakly expressed so that the head appears semiglobose in shape; frontal sculpture highly variable, ranging from smooth to sculptured throughout; female antenna with five clavomeres (defined morphologically, see Johnson 1984); notauli absent; mesoscutellar disk sculpture highly variable; T1 strongly transverse; T2 longer than wide; T2 often with distinctive coriaceous to reticulate microsculpture extending beyond the pits marking the position of the antecostal suture and the longitudinal striae arising between those pits. As Dodd (1914) noted, the ovipositor is often exserted a great distance and is easily seen, but this feature is relevant for only a minority of species. Within Phanuromyia, we separate the galeata group purely as a practical grouping, and at this point we do not assert its monophyly. The group may be distinguished, first and foremost, by their unusually large body size: most specimens are greater than 2 mm in length. Beyond that, the body is distinctly elongate, T1 in the female is produced into a horn to house the ovipositor, T2 is strongly elongate, as often are the following tergites. The group is strictly Neotropical in distribution, extending from the Isthmus of Tehuantepec in the north to Misiones in southern Paraguay. Sculpture on posterior half of mesoscutum: coriaceous to rugulose, at most with fine irregular longitudinal sculpture. Sculpture of anterior half of mesoscutellum: rugose-punctate. Thin median foliaceous lamella on propodeum: absent.

Key to assist recognition of Phanuromyia
Color of coxae: bright yellow, concolorous with legs. T1: flat, at most slightly swollen. Anterior margin of T2: with costae or foveolae throughout its width. T2 sculpture: with neither transverse series of small punctures nor scrobiculate lateral areas. Sculpture of T1: entirely costate. Posterior margin of T2: distinctly concave. Number of visible terga past T2: 2 or 3. Setation on T2: consisting of thick patches of lateral setae; consisting of widespread scattered pilosity.
Diagnosis. Phanuromyia comata can be recognized by T2 setation consisting of thick patches of lateral setae combined with widespread scattered pilosity dorsally.
Etymology. The name comata is derived from the Latin word for having long hair because this species has diagnostic patches of setae. This name is to be used as a participle.
Link to distribution map. Comments. Phanuromyia comata is the only species in the group with widespread pilosity across the entirety of T2, so this character can be used to identify a specimen very quickly. Sculpture on posterior half of mesoscutum: coriaceous to rugulose, at most with fine irregular longitudinal sculpture. Sculpture of anterior half of mesoscutellum: rugose-punctate. Thin median foliaceous lamella on propodeum: absent.

Phanuromyia constellata
Color of coxae: bright yellow, concolorous with legs. T1: flat, at most slightly swollen. Anterior margin of T2: with costae or foveolae throughout its width. T2 sculpture: with neither transverse series of small punctures nor scrobiculate lateral areas. Sculpture of T1: entirely costate. Posterior margin of T2: straight; only slightly concave. Number of visible terga past T2: 4 or 5. Setation on T2: limited to at most 1 row of setae posteriorly and sparse setation laterally.
Diagnosis. Phanuromyia corys can be recognized by the two rows of setiferous punctures converging ventrally on the frons.
Etymology. The name corys is derived from a Greek word for helmet because this species has a large head evoking the image of a knight wearing a helmet. This name is to be used as a noun in apposition.
Link to distribution map. Comments. This species is recognizable by its large size, only the largest specimens of P. odo reach over 2.7 mm in length. Phanuromyia corys may be distinguished from P. odo by the converging lines of setiferous punctures on the frons and the straight, transverse apical margin of T2.  Median keel on frons: absent. Sculpture of lower frons: with multiple transverse rugae; with irregular rugosity. Shape of mandible: slender. Median tooth of mandible: as large as adjacent teeth. Frons below median ocellus: with 2 perfectly parallel rows of setiferous punctures.
Sculpture on posterior half of mesoscutum: coriaceous to rugulose, at most with fine irregular longitudinal sculpture. Sculpture of anterior half of mesoscutellum: rugose-punctate. Thin median foliaceous lamella on propodeum: absent.
Color of coxae: bright yellow, concolorous with legs. T1: flat, at most slightly swollen. Anterior margin of T2: with costae or foveolae throughout its width. T2 sculpture: with neither transverse series of small punctures nor scrobiculate lateral areas. Sculpture of T1: entirely costate. Posterior margin of T2: straight; only slightly concave. Number of visible terga past T2: 2 or 3. Setation on T2: limited to at most 1 row of setae posteriorly and sparse setation laterally.
Diagnosis. Phanuromyia cranos can be recognized by the 2 perfectly parallel rows of setiferous punctures on the frons.
Etymology. The name cranos is derived from a Greek word for helmet because this species has a large head evoking the image of a knight wearing a helmet. This name is to be used as a noun in apposition.
Link to distribution map. Comments. Phanuromyia cranos can be identified most quickly by the distinctive pattern of setiferous punctures on its frons. The setiferous frontal puncture of P. odo may at times appear similar, but P. odo has the apical margin of T2 distinctly concave. Description. Female body length: 1.33-1.82 mm (n=20).
Sculpture on posterior half of mesoscutum: coriaceous to rugulose, at most with fine irregular longitudinal sculpture. Sculpture of anterior half of mesoscutellum: smooth. Thin median foliaceous lamella on propodeum: absent.
Color of coxae: bright yellow, concolorous with legs. T1: distinctly swollen throughout entire length. Anterior margin of T2: with costae or foveolae throughout its width. T2 sculpture: with neither transverse series of small punctures nor scrobiculate lateral areas. Sculpture of T1: almost entirely smooth. Posterior margin of T2: straight. Number of visible terga past T2: 3 or 4. Setation on T2: limited to at most 1 row of setae posteriorly and sparse setation laterally.
Diagnosis. Phanuromyia cudo can be recognized by the swelling of the entire T1 segment. Etymology. The name cudo is derived from the Latin word for a helmet made of raw skin because this species has a large head evoking the image of a knight wearing a helmet. This name is to be used as a noun in apposition.
Comments. This species exhibits variation in several characters, but can be reliably diagnosed by the swollen appearance of the entirety of T2. Sculpture on posterior half of mesoscutum: coriaceous to rugulose, at most with fine irregular longitudinal sculpture. Sculpture of anterior half of mesoscutellum: rugose-punctate. Thin median foliaceous lamella on propodeum: absent.

Phanuromyia dissidens
Color of coxae: bright yellow, concolorous with legs. T1: flat, at most slightly swollen. Anterior margin of T2: with costae or foveolae throughout its width. T2 sculpture: with neither transverse series of small punctures nor scrobiculate lateral areas. Sculpture of T1: costate at sides, smooth medially; entirely costate. Posterior margin of T2: straight; slightly convex. Number of visible terga past T2: 5. Setation on T2: limited to at most 1 row of setae posteriorly and sparse setation laterally.
Diagnosis. Phanuromyia dissidens can be recognized by the even covering of setiferous punctures on the frons.
Etymology. The name dissidens is derived from the Latin word for differing because this species has an evenly punctured frons, differentiating it from the other species in the group. This name is to be used as a participle. Sculpture on posterior half of mesoscutum: with strong, parallel longitudinal keels. Sculpture of anterior half of mesoscutellum: smooth. Thin median foliaceous lamella on propodeum: absent.
Color of coxae: dark brown to black, contrasting with legs. T1: flat, at most slightly swollen. Anterior margin of T2: with costae or foveolae throughout its width. T2 sculpture: with neither transverse series of small punctures nor scrobiculate lateral areas. Sculpture of T1: entirely costate. Posterior margin of T2: straight; slightly convex. Number of visible terga past T2: 5. Setation on T2: limited to at most 1 row of setae posteriorly and sparse setation laterally.
Diagnosis. Phanuromyia galeata can be recognized by the median keel on the frons.
Etymology. The name galeata is derived from a Latin word for helmet because this species has a large head evoking the image of a knight wearing a helmet. This name is to be used as a noun in apposition.
Link to distribution map. Comments. This species most closely resembles P. galerita, but the two can be easily distinguished from each other by comparing the mandibles: P. galeata has a slender mandible with a small median tooth, while P. galerita has much broader mandibles and a median tooth as large as the outer teeth.  Sculpture on posterior half of mesoscutum: with strong, parallel longitudinal keels. Sculpture of anterior half of mesoscutellum: rugose-punctate. Thin median foliaceous lamella on propodeum: absent.
Color of coxae: dark brown to black, contrasting with legs. T1: flat, at most slightly swollen. Anterior margin of T2: with costae or foveolae throughout its width. T2 sculpture: with neither transverse series of small punctures nor scrobiculate lateral areas. Sculpture of T1: entirely costate. Posterior margin of T2: straight; slightly convex. Number of visible terga past T2: 5. Setation on T2: limited to at most 1 row of setae posteriorly and sparse setation laterally.

Diagnosis.
Phanuromyia galerita can be recognized by the median tooth of the mandible, which is as large as the adjacent teeth.
Etymology. The name galerita is derived from the Latin word for wearing a hood because this species has a large head evoking the image of a hooded figure. This name is to be used as a noun in apposition. This name is to be used as a participle.
Link to distribution map. Comments. This species most closely resembles P. galeata, but the two can be easily distinguished from each other by comparing the mandibles: P. galeata has a slender mandible with a small median tooth, while P. galerita has much broader mandibles and a median tooth as large as the outer teeth. Median keel on frons: absent. Sculpture of lower frons: with irregular rugosity; with 3-6 transverse rugae medially. Shape of mandible: slender. Median tooth of mandible: diminished. Frons below median ocellus: with 2 rows of setiferous punctures.

Phanuromyia hjalmr
Sculpture on posterior half of mesoscutum: coriaceous to rugulose, at most with fine irregular longitudinal sculpture. Sculpture of anterior half of mesoscutellum: rugosepunctate. Thin median foliaceous lamella on propodeum: absent.
Color of coxae: bright yellow, concolorous with legs. T1: flat, at most slightly swollen. Anterior margin of T2: with costae or foveolae throughout its width. T2 sculpture: with neither transverse series of small punctures nor scrobiculate lateral areas. Sculpture of T1: evenly costate across anterior 1/3 to 1/2, smooth in remaining apical portion. Posterior margin of T2: distinctly concave. Number of visible terga past T2: 2 or 3. Setation on T2: limited to at most 1 row of setae posteriorly and sparse setation laterally.
Diagnosis. Phanuromyia hjalmr can be recognized by the sculpture of T1, which is evenly costate across the anterior 1/3 to 1/2 and smooth in remaining apical portion, combined with the distinctly concave posterior margin of T2.
Etymology. The name hjalmr is derived from the Old Norse word for helmet because this species has a large head evoking the image of a knight wearing a helmet. This name is to be used as a noun in apposition.
Link to distribution map. Comments. This species most closely resembles P. tonsura, but the two species can be distinguished by their T1 sculpture: P. tonsura has the medial portion of T1 smooth from its anterior to posterior margin, while P. hjalmr is sculptured across the entire anterior third of T1. Sculpture on posterior half of mesoscutum: coriaceous to rugulose, at most with fine irregular longitudinal sculpture. Sculpture of anterior half of mesoscutellum: rugose-punctate. Thin median foliaceous lamella on propodeum: absent.
Diagnosis. Phanuromyia krossotos can be recognized by the thick lateral patches of setae on T2.
Etymology. The name krossotos is derived from the Greek word for fringed because this species has a distinctive fringe of lateral setae on T2. This name is to be used as an adjective.
Comments. The specimens here referred to as Phanuromyia odo were originally split into two provisional species. This was based upon differences in the relative length of the wings, specifically, whether the wings reached beyond the apex of the metasoma. Figure 67 illustrates the relationship between body size and wing length: larger specimens indeed have relatively shorter wings. However, there is no distinctive gap in the ratio between the two. Therefore, we treat them here as a single species. Sculpture on posterior half of mesoscutum: coriaceous to rugulose, at most with fine irregular longitudinal sculpture. Sculpture of anterior half of mesoscutellum: rugose-punctate. Thin median foliaceous lamella on propodeum: absent.
Diagnosis. Phanuromyia pauper can be recognized by the T2 sculpture, which is scrobiculate laterally and smooth medially.
Etymology. The name pauper refers to the lack of longitudinal costae on the base of T2. This name is to be used as a noun in apposition.
Comments. This species very closely resembles P. princeps, although the two species can be distinguished easily by the sculpture of T2: P. princeps has a complete scrobiculate angled "belt" while P. pauper only has lateral costae. Figure 67. The specimens identified as Phanuromyia odo were originally split into two separate species, here labeled "small" and "large." The trend line is calculated based on data for all specimens combined. The continuity in both variables and the overlap in specimens divided a priori into small and large categories led to the conclusion that there is insufficient evidence to separate these specimens into two species. Sculpture on posterior half of mesoscutum: coriaceous to rugulose, at most with fine irregular longitudinal sculpture. Sculpture of anterior half of mesoscutellum: rugose-punctate. Thin median foliaceous lamella on propodeum: absent.
Diagnosis. Phanuromyia princeps can be recognized by the T2 sculpture, which consists of a transverse series of small punctures in the shape of an incurved chevron.
Etymology. The name princeps is derived from the prince character in the book The Prince and the Pauper by Mark Twain in reference to its similarity to P. pauper. This name is to be used as a noun in apposition.
Comments. This species very closely resembles P. pauper, although the two species can be distinguished easily by the sculpture on T2: P. princeps has a complete scrobiculate angled "belt," while P. pauper only has lateral costae. Sculpture on posterior half of mesoscutum: coriaceous to rugulose, at most with fine irregular longitudinal sculpture. Sculpture of anterior half of mesoscutellum: uncertain, smooth. Thin median foliaceous lamella on propodeum: absent.

Phanuromyia tonsura
Color of coxae: bright yellow, concolorous with legs. T1: flat, at most slightly swollen. Anterior margin of T2: with costae or foveolae throughout its width. T2 sculpture: with neither transverse series of small punctures nor scrobiculate lateral areas. Sculpture of T1: costate at sides, smooth medially. Posterior margin of T2: distinctly concave. Number of visible terga past T2: 2 or 3. Setation on T2: limited to at most 1 row of setae posteriorly and sparse setation laterally.
Diagnosis. Phanuromyia tonsura can be recognized by the sculpture of T1, which is costate laterally and smooth medially.
Etymology. The name tonsura is derived from the Latin word for a shearing and refers to the tonsure hairstyle often worn by monks, because this species has a smooth "bald" area in the middle of T1. This name is to be used as a noun in apposition.
Comments. This species most closely resembles P.
hjalmr, but the two species can be easily distinguished by their T1 sculpture: P. tonsura has the medial portion of T1 smooth from its anterior to posterior margin, while P. hjalmr is sculptured across the entire anterior third of T1. Sculpture on posterior half of mesoscutum: coriaceous to rugulose, at most with fine irregular longitudinal sculpture. Sculpture of anterior half of mesoscutellum: uncertain, rugose-punctate. Thin median foliaceous lamella on propodeum: present.

Phanuromyia tubulifer
Color of coxae: bright yellow, concolorous with legs. T1: with distinct, tubular horn reaching higher than metascutellum. Anterior margin of T2: with costae or foveolae throughout its width. T2 sculpture: with neither transverse series of small punctures nor scrobiculate lateral areas. Sculpture of T1: entirely costate. Posterior margin of T2: straight; only slightly concave. Number of visible terga past T2: 3 or 4. Setation on T2: limited to at most 1 row of setae posteriorly and sparse setation laterally.
Diagnosis. Phanuromyia tubulifer can be recognized by the distinct, tubular horn on T1 and the thin foliaceous lamella present medially on the propodeum.
Etymology. The name tubulifer refers to the presence of the diagnostic tubular horn on T1 in this species. This name is to be used as a noun in apposition.
Link to distribution map. Comments. This species is distinct due to its tubular horn, although it is unknown whether the males express this character to any significant degree. Posterior half to 2/3 of mesoscutum with strong parallel, longitudinal keels (best viewed from behind at 45-degree angle) (Figs 39, 46); all coxae dark brown to black, contrasting with the otherwise bright yellow legs (Fig. 38) ..2 -Posterior half of mesoscutum at most with fine, irregular longitudinal sculpture, usually coriaceous to rugulose (Fig. 34); all coxae bright yellow, concolorous with remaining segments of legs (Fig. 1)  T2 with a line of foveae that begins at the anterolateral margin of the sclerite and extends obliquely toward the midline (Fig. 74)  Basal ⅔ of T1 with at least a large smooth area medially, sometimes entirely smooth (Fig. 85) (Fig. 20) .............13 12(11) T1 costate throughout its length (Fig. 62) (Fig. 8); two rows of setiferous punctures below median ocellus not parallel (Fig. 12)