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Proterosceliopsidae: A new family of Platygastroidea from Cretaceous amber
expand article infoElijah J. Talamas§, Norman F. Johnson|, Chungkun Shih, Dong Ren
‡ Capital Normal University, Beijing, China
§ Florida State Collection of Arthropods, Gainesville, United States of America
| The Ohio State University, Columbus, United States of America
¶ National Museum of Natural History, Smithsonian Institution, Washington, United States of America
Open Access

Abstract

Proterosceliopsis was erected by Ortega-Blanco et al. (2014) in their treatment of scelionid genera in Cretaceous amber from Álava, Spain. The generic description appears to have been based on specimens in which only the dorsal aspects of the mesosoma and metasoma were visible, as characters of the mesopleuron, metapleuron, lateral pronotum, and ventral metasoma were not mentioned. We here provide a comprehensive description of the genus that includes characters from throughout the body and we reinterpret some of the characters presented by Ortega-Blanco et al. (2014). Our analysis of Proterosceliopsis in the context of extant and fossil platygastroids places this group as a lineage well outside of the current families. We here designate Proterosceliopsis as the type genus of a new family, Proterosceliopsidae Talamas, Johnson, Shih & Ren, fam. nov., and describe five new species: Proterosceliopsis ambulata Talamas, Shih & Ren, sp. nov., P. nigon Talamas, Shih & Ren, sp. nov., P. plurima Talamas, Shih & Ren, sp. nov., P. torquata Talamas, Shih & Ren, sp. nov., and P. wingerathi Talamas, Shih & Ren, sp. nov. We provide a key to all presently known species in the genus. The oldest known specimen of Platygastridae s.s., in Burmese amber, is presented and compared to Proterosceliopsidae fam. nov.

Keywords

fossil, morphology, Platygastridae, Scelionidae

Introduction

Platygastroid wasps are numerous in Burmese amber, comprising 16% of Hymenoptera in a recent study by Zhang et al. (2018). Some Cretaceous taxa are recognizable among the extant fauna: Platygastridae, Sparasionini and Archaeoteleia Masner (Talamas et al. 2017a). Simultaneously, many cannot be confidently placed at the family level because of the morphological disparity of the Cretaceous fauna and because the classification of Platygastroidea is in a state of flux. Zhang et al. (2018) identified the Burmese amber specimens as Scelionidae, a family that has been used to accommodate taxa that are plesiomorphic, e.g. Nixonia Masner, Archaeoteleia Masner, Sparasion Latreille (Masner 1976), or feature characters that clearly exclude them from Platygastridae, e.g. Janzenella Masner & Johnson (Masner and Johnson 2007), Plaumannion Masner & Johnson (Masner et al. 2007), Huddlestonium Polaszek & Johnson (Masner et al. 2007) and Neuroscelio Dodd (Valerio et al. 2009).

The historical concept of Scelionidae was found to be polyphyletic in the molecular analysis of Murphy et al. (2007), which Sharkey (2007) used to treat Scelionidae as a junior synonym of Platygastridae. This resolved the issue of monophyly at the family level, but in doing so transferred the problem to the subfamilies, most of which are known to be polyphyletic. There is growing consensus among analyses that Platygastridae s.s. and the “main scelionid clade” of Murphy et al. (2007) are monophyletic lineages. The latter was retrieved in an analysis of the maxillo-labial complex by Popovici et al. (2017), and in unpublished molecular phylogenies of Platygastroidea that focus on relationships within Scelionidae (Chen et al.) and Platygastridae s.s. (Blaimer et al.). Both of these efforts include basal lineages and retrieve Platygastridae s.s. and the “main scelionid clade” as monophyletic taxa. A classification that employs the historical concept of Platygastridae and a modified concept of Scelionidae are thus well supported and provide cohesion with much of the taxonomic literature. Together, these taxa comprise the vast majority of platygastroid species, and a multi-family classification system provides better framework for higher taxonomy in the superfamily. We defer changes to family classification to the final results of the aforementioned analyses and note that while affinities are clearly present, we cannot place Proterosceliopsis in any of the existing or potential suprageneric taxa without drastically and unnecessarily altering their limits. Our analysis of Proterosceliopsis is made in the context of extant specimens from a broad geographic and taxonomic sampling, fossils from Lebanese, Burmese, Baltic, and Dominican amber, and with consideration of lineages for which additional families may be erected.

For the purposes of this paper, Archaeoteleia, Neuroscelio, Huddlestonium and Plaumannion are considered incertae sedis and are referenced by their generic names; Scelionidae refers to the “main scelionid clade” of Murphy et al. (2007); Platygastridae refers to the historical, pre-Sharkey (2007) concept of the family (=Platygastrinae + Sceliotrachelinae + Orwellium Johnson, Masner & Musetti); Nixoniidae is treated as a monogeneric family (=Nixonia) sensu Mckellar and Engel (2012); and Sparasionini refers to the concept of Johnson et al. (2008a).

Criticism of previous taxonomy

Preservation artifacts complicate the taxonomic treatment of fossils. This problem is exacerbated when primary types do not display characters in sufficient detail to enable unambiguous determination at the species level. In the case of Proterosceliopsis, P. masneri Ortega-Blanco, McKellar & Engel can be reliably separated from the species here described from Burmese amber, but some of the most important generic characters are not visible in the specimens and are missing from the description presented by Ortega-Blanco et al. (2014). As a result, taxon-defining characters cannot presently be verified in the specimen that carries the name of a species, genus, and now, a family. We redefine Proterosceliopsis because we consider P. masneri and the specimens in Burmese amber to be congeneric based on the congruence of observable characters. In this case, attaching a concept to an existing name can be defended by the available data and reduces the number of superficially defined genera in Platygastroidea. We assert that dubiously delimited taxa can be avoided by a more rigorous approach to the treatment of fossils, one in which taxa are described only when based on well-preserved specimens and with the necessary background knowledge about the superfamily.

Material and methods

Informatics

The numbers prefixed with acronyms, e.g. “USNMENT” or “OSUC”, are unique identifiers for the individual specimens (note the blank space after some acronyms). Details on the data associated with these specimens may be accessed at the following link: purl.oclc.org/NET/hymenoptera/hol and entering the identifier in the form. Persistent URIs for each taxonomic concept were minted by xBio:D in accordance with best practices recommend by Hagedorn et al. (2013). Morphological terms were matched to concepts in the Hymenoptera Anatomy Ontology (Yoder et al. 2010) using the text analyzer function. A table of morphological terms and URI links is provided in Appendix 1. Taxonomic synopses and matrix-based descriptions were generated from the Hymenoptera Online Database (hol.osu.edu) and the online program vSysLab (vsyslab.osu.edu) in the format of character: state. Characters that were not visible because of preservation or orientation of the insects are coded as “not visible”.

Imaging and microscopy

The amber pieces were cut and polished to optimize the viewing and photography of specimens for taxonomic study. Direct examination of the specimens was made with a Zeiss V8 stereomicroscope and an Olympus BX51 compound microscope.

Photographs were captured with multiple imaging systems: a Z16 Leica lens with a JVC KY-F75U digital camera using Cartograph and Automontage software; an Olympus BX51 compound microscope with a Canon EOS 70D digital SLR camera; and a Leica DM2500 compound microscope with a Leica DFC425 camera. Illumination was achieved with a lighting dome or with LED gooseneck lamps and mylar light dispersers. Images were rendered from z-stacks with Automontage, Helicon Focus or Zerene Stacker. In some cases, multiple montage images were stitched together in Photoshop to produce larger images at high resolution and magnification. Full resolution images are archived at the image database at The Ohio State University (specimage.osu.edu).

Dissections for scanning electron microscopy were performed with a minuten probe and forceps and body parts were mounted to a 12 mm slotted aluminum mounting stub (EMS Cat. #75220) using a carbon adhesive tab (EMS Cat. #77825-12) and sputter coated with approximately 70 nm of gold/palladium using a Cressington 108 auto sputtercoater. Micrographs were captured using a Hitachi TM3000 Tabletop Microscope at 15 keV.

Author contributions

EJT: photography, scanning electron microscopy, taxon concepts, manuscript preparation; NFJ: manuscript preparation; CKS: manuscript preparation, provision of Burmese amber; DR: manuscript preparation, provision of Burmese amber.

Collections

The amber specimens of Proterosceliopsis studied here were collected from Kachin (Hukawng Valley) of northern Myanmar, which was dated at 98.79+0.62 Ma (Cruickshank and Ko 2003, Shi et al. 2012), equivalent to the earliest Cenomanian and approximately 1 Myr within the boundary between the Early and Late Cretaceous (Walker et al. 2012). This deposit yielded many well-preserved insect fossils (Chen et al. 2018a–b, Li et al. 2018, Wang et al. 2016, Zhang et al. 2018).

Specimens on which this work is based are deposited in the following repositories with abbreviations used in the text:

CCHH Hoffeins Collection, Hamburg, Germany

CNCI Canadian National Collection of Insects, Ottawa, Canada

CNU Key Lab of Insect Evolution and Environmental Changes, Capital Normal University, Beijing, China

FSCA Florida State Collection of Arthropods, Gainesville, FL, USA

KUNH Kansas University Natural History Museum, Lawrence, KS, USA

OPPC Ovidiu Popovici, personal collection, “A.I. Cuza” University, Faculty of Biology, Iasi, Romania

OSUC C.A. Triplehorn Insect Collection, The Ohio State University, Columbus, OH, USA

USNM National Museum of Natural History, Washington, DC, USAA

Character annotations

1Rs (Fig. 15) 1st radial sector vein

2Rs (Fig. 15) 2nd radial sector vein

b (Fig. 15) bulla

C, C+R (Fig. 15) marginal vein

Cu (Fig. 15) cubital vein

fas (Fig. 10) facial striae

ff (Figs 43–44) felt field

M (Fig. 15) medial vein

mas (Fig. 10) malar striae

mees (Figs 16, 20) mesepimeral sulcus

ms (Figs 7–8, 10) malar sulcus

net (Figs 16–17, 19) netrion

p (Fig. 22) cuticular pores

pp (Fig. 17) mesopleural pit

prcs (Figs 16, 19, 21) pronotal cervical sulcus

ps (Figs 1–6) papillary (basiconic) sensilla

r (Fig. 15) stigmal vein

R1 (Fig. 15) postmarginal vein

Rs+M (Fig. 15) basal vein

sk (Figs 11–13) skaphion

Sc+R (Fig. 15) submarginal vein

S6 (Fig. 47) metasomal sternites 6

T6–T8 (Figs 4549, 55, 61) metasomal tergites 6–8

tel (Figs 16, 19, 21) transepisternal line

Results

Character discussion

Most of the diagnostic characters of Proterosceliopsis can be found in extant platygastroids but are present in a unique combination in this genus. Additionally, Proterosceliopsis exhibits significant differences from each of the taxa with which it shares characters. We present discussions of these characters and their distribution among platygastroid taxa as a prelude to the generic treatment.

Antenna

Bin (1981) defined the antennal clava in Telenominae (Scelionidae) based on the presence of papillary sensilla on the ventral side of the distal antennomeres (previously referred to as basiconic sensilla). The presence of these sensilla on female antennae is a synapomorphy for Platygastroidea (Austin et al. 2005), and Bin’s definition of the clava has since been applied to all members of the superfamily. We continue to define the clava on the basis of papillary sensilla instead of relative antennomere size for multiple reasons: antennomeres can increase in size gradually (Figs 2–4); the size among antennomeres with papillary sensilla can be variable (Fig. 5); distinctly enlarged antennal segments may not have papillary sensilla, as in the clubbed antennae found in most males of Helava Masner & Huggert (Masner and Huggert 1989, Talamas and Masner 2016). The claval formula is often of taxonomic value, particularly at the species-level, and its use requires that specimens be preserved well enough to observe these structures. Small bubbles in the amber are sometimes visible at the tips of the sensilla (Fig. 6) and can be useful for identifying their presence.

Figures 1–6. 

1 Sceliomorpha Ashmead (DPI_FSCA 00008723), female antenna, ventral view 2 Electroteleia (DPI_FSCA 00010129), female antenna, ventral view 3 Proterosceliopsis plurima (CNU-HYM-MA-2016102), female antenna, ventral view 4 Nixonia watshami Johnson & Masner (OSUC 149432), female antenna, ventral view 5 Proterosceliopsis nigon (CNU-HYM-MA-2017566), female antenna, ventral view. 6 Proterosceliopsis wingerathi (CNU-HYM-MA-2016101), female antenna, lateral view. Scale bars in millimeters.

A transverse orientation of papillary sensilla is known only in Nixonia (Fig. 4) (Johnson and Masner 2006) whereas all other extant platygastroids have a longitudinal arrangement (Fig. 1). We found that in some species of Proterosceliopsis (Figs 3, 5) and Cretaceous Electroteleia Brues (Fig. 2) the sensilla are arranged at an oblique angle. This was found in multiple specimens in which the general shape of the antennomeres is intact, and thus we do not attribute this to taphonomic deformation. Instead, we consider that different arrangements of these sensilla were possible earlier in the history of Platygastroidea.

Malar sulcus

The malar sulcus found in Proterosceliopsis is unaccompanied by facial or malar striae (Fig. 7), a state which can be found in some Scelionidae (Fig. 8) and some Sparasionini (Electroteleia). In Platygastridae, the malar sulcus is present in only two genera: Metaclisis Förster and Orseta Masner & Huggert, each of which have both facial and malar striae (Fig. 10) (Masner and Huggert 1989), and Orwellium, which does not have facial or malar striae. The malar sulcus is entirely absent in Nixonia Masner (Fig. 9).

Figures 7–10. 

7 Proterosceliopsis plurima (CNU-HYM-MA-2016102), head, anteroventral view 8 Calliscelio Ashmead (CNC494853), head, anteroventral view 9 Nixonia watshami (OSUC 149432), head, anteroventral view 10 Metaclisis (USNMENT01029162), head, anteroventral view. Scale bars in millimeters.

Palpal formula

Ortega-Blanco et al. (2014) recorded the palpal formula of P. masner to be 5:3. We observed 5-merous, cylindrical maxillary palps in P. nigon, P. torquata, and P. wingerathi; and at least 4 maxillary palpomeres in P. plurima. Two labial palpomeres are visible in P. torquata. Five-merous palpomeres were retrieved as the plesiomorphic condition for Platygastroidea in the phylogenetic analysis by Popovici et al. (2017) and are found in Sparasionini, Nixonia, and Archaeoteleia. This indicates that Proterosceliopsis is also a basal lineage well outside of Scelionidae in which the palpal formula is 4:2 or less (Popovici et al. 2017), and Platygastridae in which the palpal formula is 2:1 or less (Popovici, personal communication).

Skaphion

The anterior mesoscutum in most species of Proterosceliopsis features a smooth, transverse structure to which we apply the term skaphion (Figs 11, 50, 58, 62). We do not assert that the skaphion of Proterosceliopsis is homologous with that found in scelionid genera (Figs 12–13) and note that it can be found in other Cretaceous taxa that do not belong to Proterosceliopsis or Scelionidae, e.g. Electroteleia (Fig. 14).

Figures 11–14. 

11 Proterosceliopsis torquata (CNU-HYM-MA-2016106), head and mesosoma, dorsal view 12 Calotelea Westwood (OSUC 56216), head and mesosoma, dorsal view 13 Nyleta striaticeps Dodd (OSUC 174452), mesosoma, anterior view 14 Electroteleia (CNU-HYM-MA-2016103), mesosoma and metasoma, dorsolateral view. Scale bars in millimeters.

Costal vein

In Proterosceliopsis, the costal vein is present anterior to the fusion with R and extends proximally beyond the bulla (Fig. 15). This form is known from other Cretaceous platygastroids (Fig. 14), but not in extant taxa.

Figure 15. 

Proterosceliopsis torquata (CNU-HYM-MA-2016106), fore wing, dorsal view. Scale bar in millimeters.

Setation of the pronotal cervical sulcus

The pronotal cervical sulcus can take many forms, including a distinct line of foveae (Fig. 18), a well-defined smooth groove (Fig. 19), or a weakly defined furrow along the lateral pronotal rim (Fig. 17). In Nixonia, Johnson and Masner (2006) reported an area of dense setation along the anterior margin of the lateral pronotum that is often associated with solidified exudate (Fig. 20). Setation along the pronotal cervical sulcus is found in Proterosceliopsis (Fig. 16, 52, 59) and many platygastrids (Figs 19, 26), and often contains what appears to be solidified exudate. The unpublished phylogenetic analysis of Platygastridae (Blaimer et al.) indicates that stem lineages of this family have a setose pronotal cervical sulcus. Figures 65–66 illustrate the oldest known platygastrid, in Burmese amber, that exhibits this character, consistent with our treatment of it as a plesiomorphy for Platygastridae that varies significantly in some derived platygastrid lineages (Figs 21–26). Figures 21–22 illustrate the unusual form found in Sacespalus in which setation is absent and the pronotal cervical sulcus is broadly expanded. Pores can be seen in the ventral portion of the pronotal cervical sulcus of Sacespalus (Fig. 22), which is congruent with the hypothesis that this character is associated with glandular secretions. We do not know of any scelionids with a setal patch along the anterior portion of the lateral pronotum but note that the antespiracular setal patch (sensu Yoder et al. 2012) in Scelionidae is associated with cuticular pores (Fig. 28). Based on the pronotal position of these setal patches, we suspect that they have similar function.

Figures 16–20. 

16 Proterosceliopsis plurima (CNU-HYM-MA-2016102), mesosoma, lateral view 17 Scelio Latreille (USNMENT00989612_3), mesosoma, ventrolateral view 18 Archaeoteleia gracilis Masner (OSUC 163002), mesosoma, ventrolateral view 19 Fidiobia (USNMENT01197212_2), mesosoma, ventrolateral view 20 Nixonia watshami (OSUC 149432), head, lateral view. Scale bars in millimeters.

Figures 21–26. 

21 Sacespalus Kieffer (USNMENT01197981_1), mesosoma, lateral view 22 Sacespalus (USNMENT01197981_1), ventral portion of pronotal cervical sulcus, lateral view 23 Leptacis Förster (USNMENT00872705), anterior portion of pronotal cervical sulcus, lateral view 24 Synopeas Förster (USNMENT00989616_3), mesosoma, lateral view 25 Platygaster Latreille (USNMENT01197214_1), pronotum, lateral view 26 Trichacis Förster (USNMENT00989620), pronotum, lateral view. Scale bars in millimeters.

Figures 27, 28. 

27 Doddiella (OSUC 56323), head and mesosoma, lateral view 28 Doddiella (USNMENT00872797), pronotum and mesopleuron, lateral view. Scale bars in millimeters.

Netrion

A netrion is clearly present in 3 of the 5 species that we here describe. Mikó et al. (2007) reported that in Nixonia the trachea associated with the anterior thoracic spiracle extends ventrally between the netrion sulcus and the posterior pronotal inflection. Externally this results in the netrion sulcus dorsally terminating anterior to the anterior thoracic spiracle (Fig. 20). This character can be used to separate Nixonia from nearly all other platygastroids, in which the netrion sulcus, when present, terminates posterior or ventral to the anterior thoracic spiracle (Figs 16–18, 20).

Transepisternal line

Among extant platygastroids, the transepisternal line is found exclusively in Platygastridae (Fig. 19). We treat it as a plesiomorphy for Platygastridae based on its presence in the Cretaceous specimen illustrated in Figs 65–66, and because it is found in stem lineages of the family based on a preliminary analysis of molecular data (Blaimer et al.). The internal anatomy associated with this structure has yet to be examined in detail and will likely shed light on its function and evolution within Platygastridae. The transepisternal line in Proterosceliopsis is clearly present in all specimens from Burmese amber (Figs 16, 53) suggesting a close relationship between Proterosceliopsidae and Platygastridae.

Mesepimeral sulcus

The presence of a fully developed mesepimeral ridge was retrieved by Vilhelmsen et al. (2010) as a potential autapomorpy for Proctotrupomorpha s.s. This ridge corresponds externally to the mesepimeral sulcus, which is found in all platygastroid families except Platygastridae (Figs 16–20). We consider the loss of the mesepimeral sulcus to be an apomorphy for Platygastridae, but whether this internally corresponds to loss of the mesepimeral ridge has yet to be investigated. Some examples of an absent or weakly indicated mesepimeral sulcus can be found in Scelionidae. In some cases, these are clearly secondary derivations that occurred at the species level (Fig. 17). In the unusual Doddiella Kieffer (Scelionidae) the mesepimeral sulcus is not indicated externally but the mesepimeral ridge can be seen through the semitransparent exoskeleton (Fig. 27). The presence of the mesepimeral sulcus in Proterosceliopsidae provides a reliable mesosomal character to separate it from Platygastridae.

Setation and sulci of the metasoma

Proterosceliopsis from Burmese amber and Nixonia share the presence of transverse, depressions along the anterior margins of T1–T5 (Figs 29, 32) and S1–S5 (Figs 34, 37). In Nixonia, these depressions have visible setation. In Proterosceliopsis they appear to have very fine setation, but the cloudy exudate prevents an unobscured view. In P. torquata these depressions are not clearly visible on T6, and in P. plurima they are not clearly present on S6. In the other Burmese species of Proterosceliopsis, and in all species of Nixonia, these depressions are present on T6 and S6. We suspect that they are present on T6 and S6 in all species of Proterosceliopsis, but the preservation of these specimens prevents us from reaching a confident conclusion.

Figures 29–33. 

29 Proterosceliopsis nigon (CNU-HYM-MA-2017566), head, mesosoma, metasoma, dorsal view 30 Trimorus Förster (USNMENT01197861), metasoma, dorsal view 31 Amitus Haldeman (OSUC 404941), metasoma, dorsal view 32 Nixonia krombeini Johnson & Masner (OSUC 146429), metasoma, dorsal view 33 Sparasion philippinensis Kieffer (USNMENT00872835), metasoma, dorsal view. Scale bars in millimeters.

The two components of this character, setation/glandular secretion and sulci along the anterior tergites and sternites, vary independently among platygastroid families. Sparasionini and Archaeoteleia feature transverse sulci across the anterior margins of the external tergites and sternites (Figs 33, 38, 44), which may sometimes be reduced on T6 and S6 in females of Electroteleia (Sparasionini) and in Archaeoteleia. These sulci are found in all specimens that we examined in Lebanese amber, which constitute the oldest platygastroid fossils, and we consider it likely that this is the plesiomorphic condition for Platygastroidea. The metasoma in Sparasionini and Archaeoteleia lacks setation on the anterior tergites and sternites beyond T1 and S1, but there may be numerous felt fields (Fig. 44).

Figures 34–38. 

34 Proterosceliopsis nigon (CNU-HYM-MA-2017566), head, mesosoma, metasoma, ventral view 35 Dvivarnus mikuki Talamas & Mikó (USNMENT01059135), metasoma, ventral view 36 Trichacis (USNMENT01059347), metasoma, ventral view 37 Nixonia watshami (OSUC 149432), metasoma, ventral view 38 Sparasion philippinensis (USNMENT00872835), metasoma, ventral view. Scale bars in millimeters.

Figures 39–44. 

39 Trichoteleia bidentata Talamas (CASENT 2132802), T1, lateral view 40 Scelioninae (OSUC 254572), S1-S2, ventrolateral view 41 Neuroscelio doddi (OSUC 147252), S1-S2, ventrolateral view 42 Dvivarnus mikuki (USNMENT01059135), T3-T5, dorsal view 43 Helava alticola Masner & Huggert (USNMENT00989211), metasoma, lateral view 44 Archaeoteleia gracilis (OSUC 163002), metasoma, lateral view. Scale bars in millimeters.

In Platygastridae and Scelionidae, the anterior portions of T1–T2 and S1–S2 typically feature a transverse line of foveae or pits, which may or may not be clearly defined (Figs 30–31, 35–36). The transverse sulci in scelionids are not associated with setation, and patches of dense setae tend to be located on the lateral portion of the tergite, sometimes in well-defined pits (Figs 39–40). Some teleasines (Scelionidae) exhibit a transverse sulcus along anterior T3 and S3 (Fig. 30, 35) and setal patches with pores on T4–T5 (Fig. 42). Platygastridae exhibits the greatest variation in the size, location, and shape of setal pits on metasomal segments 1 and 2. They may be present laterally or medially, as a broad transverse patch that spans the width of the tergite or sternite (Fig. 43), in well-defined pits (Fig. 31), or they may be absent entirely (e.g. Orwellium Johnson, Masner & Musetti). Felt fields, which are absent in Nixonia and Proterosceliopsis, are found only on S2 in Platygastridae and Scelionidae, with a few exceptions (e.g. Heptascelio Kieffer (Johnson et al. 2008b)). Neuroscelio doddi Galloway, Austin & Masner exhibits an unusual and noteworthy form of sternal setation in which dense tufts are located medially on S1–S2 (Fig. 41).

Setal patches on the metasoma and pronotum, which are often associated with pores in the integument, clearly warrant further examination. A survey of platygastroid morphology with a scanning electron microscope has revealed an array of additional pore locations and forms on the head and mesosoma. Our present analysis of these characters, which provides only a broad overview as it relates to family level classification of Proterosceliopsis, will be developed further in future projects. The presence of cuticular pores and solidified exudate in the same locations point to the activity of internal glands.

Ovipositor system

Ortega-Blanco et al. (2014) interpreted T7+8 to be the ovipositor, which they described as short and broad. We here clarify that these sclerites are not the ovipositor, but they could be considered part of the ovipositor system sensu Austin and Field (1997). The known diversity of ovipositor systems in Platygastroidea has expanded in recent years, with at least two derivations of a telescoping ovipositor system found in Platygastridae (Talamas et al. 2017b). Extension of the ovipositor system in Proterosceliopsis appears to operate via telescoping membrane between T6 and T7, with T7 and T8 clearly present as separate sclerites in some specimens (Fig. 45). The unpublished phylogenetic analyses suggest that conjunctival expansion between T6 and T7 occurred independently in Scelionidae and Archaeoteleia. The placement of Proterosceliopsis well outside of these families indicates that expansion of conjunctiva between T6 and T7 into a telescoping ovipositor system has occurred at least three times: in Proterosceliopsis, Archaeoteleia, and Scelionidae. We also have found evidence of conjunctival expansion between T6 and T7 in the platygastrid genus Fidiobia Ashmead. It does not appear to be telescoping but provides clear evidence of yet another independently derived elongation of conjunctiva between T6 and T7 (Figs 46–47), lending evidence to the plasticity of this morphological system.

Figures 45–47. 

45 Proterosceliopsis ambulata (CNU-HYM-MA-2016105), distal metasoma, dorsal view 46 Fidiobia (USNMENT01197212_2), distal metasoma, dorsal view 47 Fidiobia (USNMENT01197212_2), distal metasoma, ventrolateral view.

Taxonomy

Proterosceliopsidae Talamas, Johnson, Shih & Ren, fam. nov.

Diagnosis

Antenna with 14 or 15 antennomeres; malar sulcus present (Fig. 7); facial striae absent (Fig. 7); malar striae absent (Fig. 7); pronotal cervical sulcus present as a furrow of fine setation associated with glandular excretion (Figs 16, 48–49, 5153, 59); mesopleuron with transepisternal line and mesepimeral sulcus (Figs 16, 5153, 59, 63); T3–T5 and S3–S5 anteriorly with depressions associated with glandular excretion (Figs 29, 34, 56, 58, 62).

Proterosceliopsis Ortega-Blanco, McKellar & Engel

Proterosceliopsis Ortega-Blanco, McKellar & Engel, 2014: 554 (original description. Type: Proterosceliopsis masneri Ortega-Blanco, McKellar & Engel, by monotypy and original designation. Diagnosis); Talamas et al. 2017a: 251, 253 (description, keyed).

Diagnosis

See family diagnosis.

Description

Head: Facial striae: absent. Malar sulcus: present. Malar striae: absent. Orbital carina: absent. Setation of compound eye: absent. Torulus: opening anteriorly. Frontal ledge: absent. OOL: lateral ocellus separated from compound eye by less than one ocellar diameter. Macrosculpture of head: absent. Hyperoccipital carina: absent. Occipital carina: present, continuous dorsally and ventrally extending to posterior articulation of the mandible.

Mesosoma: Propleural epicoxal sulcus: absent. Posterolateral corner of propleuron: strongly pointed. Pronotal cervical sulcus: furrow of dense fine setae. Epomial carina: absent. Transverse pronotal carina: absent. Antero-admedian lines: absent. Macrosculpture of mesosoma: absent. Orientation of notauli: converging posteriorly. Mesoscutal humeral sulcus: indicated by smooth furrow. Posterior mesoscutellar sulcus: foveate. Sculpture of metanotal trough: foveate. Metascutellum: undifferentiated from metanotal trough. Ventral mesopleural furrow: present. Ventral mesopleural carina: present. Mesopleural carina: absent. Anterior mesepisternal area: absent. Episternal foveae: absent. Transepisternal line: present. Mesopleural pit: absent. Prespecular sulcus: absent. Mesepimeral sulcus: present. Metapleural carina: present. Ventral metapleural area: convex and without macrosculpture. Dorsal metapleural area: convex and without macrosculpture. Ventral surface of metapleuron: setose. Sculpture of dorsal propodeum: coarsely rugose. Setal patch on anterodorsal surface of hind coxa: present.

Fore Wing: submarginal (Sc+R), marginal (C+R), postmarginal (R1) and stigmal vein (r) present; C extending proximally past bulla; 1Rs short and nebulous; 2Rs sclerotized and extending to wing margin. Basal vein (Rs+M) and M+Cu nebulous to weakly sclerotized. Median vein (M) present as a nebulous line in distal portion of the wing; tibial spur formula: 1-2-2.

Metasoma: T1–T5 with depressions anteriorly (unclear in P. masneri); S1–S5 with depressions anteriorly; 6–7 visible tergites when ovipositor not extruded. T7+T8 extruded with ovipositor system (based on P. ambulata).

Distribution

The presence of Proterosceliopsis in Burmese and Álava (Spain) amber indicates that this was a widespread genus.

Comments

Our generic description of Proterosceliopsis is based largely on specimens from Burmese amber, which provide far greater detail than the original description, and the two descriptions are congruent as far as can be observed. However, it should be noted that depressions along the anterior margin of the metasomal tergites in Proterosceliopsis masneri are not mentioned or illustrated in Ortega-Blanco et al. (2014). In their photographs of the dorsal habitus (Fig. 6A–B) it is unclear if these depressions are present. In cases where exudate is absent, photography of these depressions can be difficult, particularly in darkly colored specimens and when the amber is turbid.

Key to species of Proterosceliopsis (females)

1 Antenna with 15 antennomeres P. plurima Talamas, Shih & Ren, sp. nov.
Antenna with 14 antennomeres 2
2 T1 with horn (Fig. 58) 3
T1 without horn (Figs 29, 48, 62) 4
3 T6 distinctly the longest tergite (Fig. 58) P. torquata Talamas, Shih & Ren, sp. nov.
T1–T6 approximately equal in length P. masneri Ortega-Blanco, McKellar & Engel
4 T1 evenly convex and without macrosculpture or raised area (Figs 48, 50); antenna with 7 clavomeres; A7 distinctly transverse (Fig. 49) P. ambulata Talamas, Shih & Ren, sp. nov.
T1 medially with raised area of longitudinal striation (Fig. 29); antenna with more than 7 clavomeres; A7 about as long as wide (Figs 5–6) 5
5 Antennae with 9 clavomeres (Fig. 5); T2 without striae posterior to transverse sulcus (Fig. 29) P. nigon Talamas, Shih & Ren, sp. nov.
Antennae with 8 clavomeres (Fig. 6); anteromedial T2 with longitudinal striae posterior to transverse sulcus (Fig. 62) P. wingerathi Talamas, Shih & Ren, sp. nov.

Proterosceliopsis ambulata Talamas, Shih & Ren, sp. nov.

Figures 45, 48–50

Diagnosis

Proteroscelio ambulata shares with P. wingerathi, P. nigon and P. masneri the roughly equal lengths of metasomal segments 1–6. It can be separated from all of these by the evenly convex form of T1, which has a horn in P. masneri and an anteromedian area of prominent striae in P. nigon and P. wingerathi.

Description

Head: Number of antennomeres in female: 14. Number of clavomeres in female: 7. Claval formula in female: 1-2-2-2-2-2-2. Number of mandibular teeth: 3. Number of labial palpomeres: not visible. Number of maxillary palpomeres: not visible. Shape of clypeus: narrow, transverse. Central keel: absent. Antennal scrobe: undifferentiated sculpturally from remainder of frons. Anterior margin of occipital carina: simple.

Figures 48–50. 

Proterosceliopsis ambulata (CNU-HYM-MA-2016105) 48 habitus, lateral view (left side) 49 habitus, lateral view (right side) 50 head, mesosoma, metasoma, dorsal view. Scale bars in millimeters.

Figures 51, 52. 

Proterosceliopsis plurima (CNU-HYM-MA-2016102) 51 habitus, lateral view (right side) 52 habitus, lateral view (left side). Scale bars in millimeters.

Mesosoma: Pronotal prespiracular depression: present, without striation. Netrion: absent. Skaphion: present. Posterior notaulus: reaching posterior margin of mesoscutum. Width of notaulus: expanding posteriorly. Parapsidal lines: absent. Mesoscutal suprahumeral sulcus: indicated by smooth furrow. Scutoscutellar sulcus: simple. Postacetabular carina: absent. Postacetabular sulcus: absent. Mesopleural epicoxal sulcus: present. Episternal foveae: absent. Metapleural sulcus: present as a transverse furrow. Lateral propodeal carina: present laterally as two small posteriorly-pointing projections.

Metasoma: Horn on T1 in female: absent. Sculpture of T1: weakly longitudinally striate throughout. Macrosculpture of T2–T5: absent. Anterior tergal depressions: visible on T1–T6. Median keel on S2: absent. Macroculpture of S3–S6: absent. Anterior sternal depressions: visible on S1–S6.

Etymology

This species is given the name “ambulata” because the holotype specimen appears to be walking.

Link to distribution map

Material examined

Holotype female: MYANMAR: CNU-HYM-MA-2016105 (deposited in CNU).

Proterosceliopsis masneri Ortega-Blanco, McKellar & Engel

Proterosceliopsis masneri Ortega-Blanco, McKellar & Engel, 2014: 555, 568 (original description, diagnosis, keyed).

Comments

Proterosceliopsis masneri may eventually be considered a nomen dubium as a species due to the paucity of detailed information on its morphology. The study of additional material from Álava amber is needed to clarify characters on this species, particularly the lateral mesosoma and structures on the anterior portions of the metasomal segments.

Proterosceliopsis nigon Talamas, Shih & Ren, sp. nov.

Figures 5, 29, 34

Diagnosis

Protersoceliopsis nigon is most similar to P. wingerathi, from which it can be separated by having 9 clavomeres and T2 without striation posterior to the transverse anterior depressions.

Description

Head: Number of antennomeres in female: 14. Number of clavomeres in female: 9. Claval formula in female: 1-2-2-2-2-2-2-2-2. Number of mandibular teeth: not visible. Number of labial palpomeres: not visible. Number of maxillary palpomeres: at least 5. Shape of clypeus: narrow, transverse. Central keel: absent. Antennal scrobe: indicated by faint transverse rugae. Anterior margin of occipital carina: crenulate.

Mesosoma: Pronotal prespiracular depression: present, without striation. Netrion: absent. Skaphion: present. Posterior notaulus: not reaching posterior margin of mesoscutum. Width of notaulus: expanding posteriorly. Parapsidal lines: present. Mesoscutal suprahumeral sulcus: simple, without furrow or cells. Scutoscutellar sulcus: crenulate. Postacetabular carina: present directly posterior to acetabulum. Postacetabular sulcus: present as simple furrow. Mesopleural epicoxal sulcus: present. Metapleural sulcus: present as a transverse furrow. Lateral propodeal carina: present and continuous dorsally, forming lamella surrounding metasomal depression, medial portion raised and projecting dorsally. Metasomal depression: excavate, interior surface with striae dorsomedially.

Metasoma: Horn on T1 in female: absent. Sculpture of T1: longitudinally striate medially. Macrosculpture of T2–T5: absent. Anterior tergal depressions: visible on T1–T6. Median keel on S2: absent. Macroculpture of S3–S6: longitudinal median carina on S3–S4, otherwise absent. Anterior sternal depressions: visible on S1–S6.

Etymology

This word “nigon” is Anglo-Saxon for “nine”, referring to the number of clavomeres in this species, and is treated as a noun in apposition.

Link to distribution map

Material examined

Holotype female: MYANMAR: CNU-HYM-MA-2017566 (deposited in CNU). Other material: (1 female) MYANMAR: OPPC1718 (deposited in OPPC).

Proterosceliopsis plurima Talamas, Shih & Ren, sp. nov.

Figures 3, 7, 16, 51–52, 53–55, 56–57

Diagnosis

The number of antennomeres in this species, 15, provides a simple means of separating it from other members of the genus. In P. plurima and P. torquata the 6th metasoma tergites and sternites are distinctly the longest. In the absence of antennal characters, these species can be separated on the form of the notaulus, which expands in width posteriorly in P. plurima and is of uniform width in P. torquata.

Description

Head: Number of antennomeres in female: 15. Number of clavomeres in female: 9. Claval formula in female: 1-2-2-2-2-2-2-2-1. Number of mandibular teeth: 3 on right mandible, 2 on left mandible. Number of labial palpomeres: not visible. Number of maxillary palpomeres: 4. Shape of clypeus: narrow, transverse. Central keel: present. Antennal scrobe: indicated by transverse rugae. Anterior margin of occipital carina: crenulate.

Figures 53–55. 

Proterosceliopsis plurima (CNU-HYM-MA-2016102) 53 head, mesosoma, anterior metasoma, lateral view (right side) 54 posterodorsal pronotum, lateral view 55 distal segments of metasoma, lateral view. Scale bars in millimeters.

Figures 56, 57. 

Proterosceliopsis plurima (CNU-HYM-MA-2016102) 56 habitus, dorsal view 57 head and mesosoma, ventral view. Scale bars in millimeters.

Mesosoma: Pronotal prespiracular depression: present, striate. Netrion: present. Skaphion: absent. Posterior notaulus: not reaching posterior margin of mesoscutum. Width of notaulus: expanding posteriorly. Parapsidal lines: absent. Mesoscutal suprahumeral sulcus: indicated by short line of cells. Scutoscutellar sulcus: simple. Postacetabular carina: present as short ridge laterally, carinae not meeting medially. Postacetabular sulcus: present as simple furrow. Mesopleural epicoxal sulcus: present. Episternal foveae: absent. Metapleural sulcus: present, anterodorsal portion comprised of cells. Lateral propodeal carina: present and continuous dorsally, forming lamella surrounding metasomal depression.

Metasoma: Horn on T1 in female: absent. Sculpture of T1: longitudinally striate medially. Macrosculpture of T2–T5: longitudinal median carina on T2 and T3, otherwise absent. Anterior tergal depressions: visible on T1–T6. Median keel on S2: present. Macroculpture of S3–S6: longitudinal median carina on S3–S4, otherwise absent. Anterior sternal depressions: visible on S1–S5.

Etymology

The epithet “plurima”, meaning “abundant” or “numerous”, refers to the number of antennomeres in this species, which is the greatest known in Platygastroidea.

Link to distribution map

Material examined

Holotype female: MYANMAR: CNU-HYM-MA-2016102 (deposited in CNU).

Comments

The number of clavomeres is here coded as 9, a basiconic sensillum on A15 is visible, as are paired sensilla on A14–A8

Proterosceliopsis torquata Talamas, Shih & Ren, sp. nov.

Figures 11, 15, 58–59, 60–61

Diagnosis

Proterosceliopsis torquata is most similar to P. plurima, with which it shares an elongate habitus, and having the 6th metasomal segment distinctly the longest. These can be separated by the presence of the horn on T1 in P. torquata (absent in P. plurima); the posteriorly expanded notauli in P. plurima (of uniform width in P. torquata), and by the number of antennomeres: 14 in P. torquata and 15 in P. plurima.

Description

Head: Number of antennomeres in female: 14. Number of clavomeres in female: 8. Claval formula in female: 1-2-2-2-2-2-2-1. Number of mandibular teeth: not visible. Number of labial palpomeres: at least 2. Number of maxillary palpomeres: at least 5. Shape of clypeus: not visible. Central keel: present. Antennal scrobe: indicated by transverse rugae. Anterior margin of occipital carina: crenulate.

Figures 58, 59. 

Proterosceliopsis torquata (CNU-HYM-MA-2016106) 58 habitus, dorsolateral view 59 head and mesosoma, lateral view. Scale bars in millimeters.

Figures 60, 61. 

Proterosceliopsis torquata (CNU-HYM-MA-2016106) 60 habitus, ventrolateral view 61 distal segments of metasoma, dorsal view. Scale bars in millimeters.

Mesosoma: Pronotal prespiracular depression: present, without striation. Netrion: present. Skaphion: present. Posterior notaulus: not reaching posterior margin of mesoscutum. Width of notaulus: uniform. Parapsidal lines: present. Mesoscutal suprahumeral sulcus: indicated by smooth furrow. Scutoscutellar sulcus: simple. Postacetabular carina: present as short ridge laterally, carinae not meeting medially. Postacetabular sulcus: present as simple furrow. Mesopleural epicoxal sulcus: present. Episternal foveae: absent. Metapleural sulcus: present as a transverse furrow. Lateral propodeal carina: present and continuous dorsally, forming lamella surrounding metasomal depression. Metasomal depression: deeply excavate, interior surface smooth.

Metasoma: Horn on T1 in female: present. Sculpture of T1: longitudinally striate medially. Macrosculpture of T2–T5: longitudinal median carina on anterior T2 and T3, otherwise absent. Anterior tergal depressions: visible on T1–T5. Median keel on S2: present. Macroculpture of S3–S6: absent. Anterior sternal depressions: visible on S1–S6.

Etymology

The Latin adjectival epithet “torquata”, meaning “adorned with a necklace or collar”, is given to this species for the collar-like shape of the pronotum in dorsal view.

Link to distribution map

Material examined

Holotype female: MYANMAR: CNU-HYM-MA-2016106 (deposited in CNU). Other material: (1 female) MYANMAR: OPPC1801 (deposited in OPPC).

Proterosceliopsis wingerathi Talamas, Shih & Ren, sp. nov.

Figures 6, 62–64

Diagnosis

Proterosceliopsis wingerathi is most similar to P. nigon, from which it can be separated by having eight clavomeres and the presence of longitudinal striation in the anteromedial portion of T2.

Description

Head: Number of antennomeres in female: 14. Number of clavomeres in female: 8. Claval formula in female: 1-2-2-2-2-2-2-1. Number of mandibular teeth: 3 on right mandible. Number of labial palpomeres: not visible. Number of maxillary palpomeres: at least 5. Shape of clypeus: narrow, transverse, medially concave. Central keel: absent. Antennal scrobe: undifferentiated sculpturally from remainder of frons. Anterior margin of occipital carina: crenulate.

Figures 62–64. 

Proterosceliopsis wingerathi (CNU-HYM-MA-2016101) 62 habitus, dorsal view 63 habitus, lateral view 64 habitus, ventrolateral view. Scale bars in millimeters.

Mesosoma: Pronotal prespiracular depression: present, without striation. Netrion: present. Skaphion: present. Posterior notaulus: not reaching posterior margin of mesoscutum. Width of notaulus: uniform. Parapsidal lines: present. Mesoscutal suprahumeral sulcus: indicated by smooth furrow. Scutoscutellar sulcus: crenulate. Postacetabular carina: absent. Postacetabular sulcus: absent. Episternal foveae: absent. Metapleural sulcus: present as a transverse furrow. Lateral propodeal carina: present and continuous dorsally, forming lamella surrounding metasomal depression. Metasomal depression: excavate, interior surface with striae dorsomedially.

Metasoma: Horn on T1 in female: absent. Sculpture of T1: longitudinally striate medially. Macrosculpture of T2–T5: anteromedial T2 longitudinally striate, otherwise absent. Anterior tergal depressions: visible on T1–T6. Median keel on S2: absent. Macroculpture of S3–S6: absent. Anterior sternal depressions: visible on S1–S6.

Etymology

This species is named for Jonathan Wingerath, Deputy Collections Manager for Paleobotany at the National Museum of Natural History, Washington, DC, to express our thanks for contributing his time and skills in preparing amber specimens for this and other projects.

Link to distribution map

Material examined

Holotype female: MYANMAR: CNU-HYM-MA-2016101 (deposited in CNU).

Comments on Cretaceous Platygastridae

The specimen illustrated in Figures 65–66 is the oldest representative of Platygastridae known to us. It complies with the current and historical concepts of the family: 10-merous antennae; T2 as the largest tergite; malar sulcus absent; pronotal cervical sulcus is a furrow with what appears to be solidified exudate; transepisternal line present (not clear in photographs); S1 and anterior S2 with setal patches. Perhaps most interesting, this specimen has marginal, stigmal and (short) postmarginal veins, as in Orwellium enigmaticum Johnson, Masner & Musetti, which Johnson et al. (2009) considered to be the sister to the rest of Platygastridae. This specimen is not sufficiently well preserved to be described at the species level, but we consider it relevant to this paper because it supports our contention that the transepisternal line and glandular nature of the pronotal cervical sulcus are plesiomorphies for Platygastridae.

Figures 65, 66. 

Platygastridae, female (KU-NHM-ENT Bu-007) 65 head, mesosoma, metasoma, dorsolateral view 66 head, mesosoma, metasoma, ventrolateral view. Insets are portions of image that have had the color altered to emphasize wing venation. Scale bars in millimeters.

Acknowledgements

We are grateful to Jonathan Wingerath (USNM), Minyue Ren, Huijia Cao, and Bingyu Zheng (CNU) for cutting and polishing amber. Lubomír Masner (CNCI) provided insightful discussion. Michael Engel generously provided a loan of Burmese amber to EJT, including platygastrid specimen KU-NHM-ENT Bu-007, and facilitated examination of Lebanese amber. Former Smithsonian interns Dylan Johnston-Jordan, Melanie Anderson, Luke Kresslein, Anthony Cuminale, Colin Schwantes and Cate Paxton contributed many of the scanning electron micrographs. Ovidiu Popovici (“A.I. Cuza” University, Faculty of Biology, Iasi, Romania) provided images that enabled species determination of two female specimens in his collection. We thank the Florida Department of Agriculture and Consumer Services – Division of Plant Industry for their support on this contribution. This research was supported by the National Natural Science Foundation of China (No. 31730087 and 31672323), the Program for Changjiang Scholars and Innovative Research Team in University (grant number IRT-17R75), and Project of High-level Teachers in Beijing Municipal Universities (grant number IDHT20180518).

References

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Appendix 1

A table of morphological terms and URI links.

Term Concept URI Preferred Term
T8 The tergum that is located on abdominal segment 8. http://purl.obolibrary.org/obo/HAO_0000061 abdominal tergum 8
acetabulum The scrobe that is located anteroventrally on the mesopectus and accommodates the procoxa. http://purl.obolibrary.org/obo/HAO_0000294 epicnemium
antenna The appendage that is composed of ringlike sclerites and the anatomical structures encircled by these sclerites and that is articulated with the cranium. http://purl.obolibrary.org/obo/HAO_0000101 antenna
antennal scrobe The scrobe that is located dorsally of the antennal foramen and is for the reception of the antenna. http://purl.obolibrary.org/obo/HAO_0001432 antennal scrobe
antennomere The anatomical structure that is delimited by the proximal and distal margins of the antennal sclerite. http://purl.obolibrary.org/obo/HAO_0000107 antennomere
anterior mesepisternal area The area delimited anteriorly by the pronotal-mesopectal suture, posterodorsally by the ventral limit of the subacropleural sulcus, and posteroventrally by the episternal foveal line. http://purl.obolibrary.org/obo/HAO_0002459 anterior mesepisternal area
anterior thoracic spiracle The spiracle that is located on the border of the pronotum and mesopleuron. http://purl.obolibrary.org/obo/HAO_0000582 anterior thoracic spiracle
antespiracular setal patch The setiferous patch that is located just anteriorly of the anterior thoracic spiracle. http://purl.obolibrary.org/obo/HAO_0001985 antespiracular setal patch
basal vein The wing vein that is basally located on the fore wing and connects or nearly connects the submarginal vein and the medial plus cubital veins. http://purl.obolibrary.org/obo/HAO_0000170 basal vein
bulla The anatomical region of the wing vein that is less melanized and more flexible than surrounding anatomical wing vein regions and corresponds to the intersection of the wing vein with a flexion line. http://purl.obolibrary.org/obo/HAO_0000184 bulla
carina The process that is elongate and external. http://purl.obolibrary.org/obo/HAO_0000188 carina
central keel The frontal line that is a carina. http://purl.obolibrary.org/obo/HAO_0001929 central keel
clava The anatomical cluster that is composed of apical flagellomeres bearing multiporous plates in female organism. http://purl.obolibrary.org/obo/HAO_0000203 clava
clypeus The area that corresponds to the site of origin of the clypeo-epipharyngeal muscle. http://purl.obolibrary.org/obo/HAO_0000212 clypeus
conjunctiva The area of the cuticle that is weakly sclerotized, with thin exocuticle. http://purl.obolibrary.org/obo/HAO_0000221 conjunctiva
costa The wing vein that is anterior to the subcosta and is connected to the humeral plate. http://purl.obolibrary.org/obo/HAO_0000225 costa
coxa The leg segment that is connected to the body and to the trochanter via conjunctivae and muscles. http://purl.obolibrary.org/obo/HAO_0000228 coxa
cubital vein The longitudinal vein that is posterior to the marginal vein. http://purl.obolibrary.org/obo/HAO_0000237 cubital vein
dorsal metapleural area The area that is delimited posterodorsally by the metapleural carina and anteroventrally by the metapleural sulcus. http://purl.obolibrary.org/obo/HAO_0000261 dorsal metapleural area
egg . http://purl.obolibrary.org/obo/HAO_0000286 egg
episternal foveae The row of impressions that is located on the anteroventral edge of the mesopleuron and is correspond to the site of origin of the mesopleuro-mesobasalar muscle. http://purl.obolibrary.org/obo/HAO_0001509 episternal foveae
eye The compound organ that is composed of ommatidia. http://purl.obolibrary.org/obo/HAO_0000217 eye
facial striae The anatomical cluster anterior to the malar sulcus that is composed of carinae radiating from the pleurostomal condyle. http://purl.obolibrary.org/obo/HAO_0002376 facial striae
felt field The setiferous patch that is located sublaterally on an abdominal sternum. http://purl.obolibrary.org/obo/HAO_0000322 felt field
fore wing The wing that is located on the mesothorax. http://purl.obolibrary.org/obo/HAO_0000351 fore wing
frons The area that is located dorsally of the ventral margin of the antennal rim and ventrally of the anterior ocellus medial to the inner margins of the eye and malar line. http://purl.obolibrary.org/obo/HAO_0001044 upper face
frontal ledge The edge that traverses the upper face between the antennal foramen and the median ocellus separating a more horizontal dorsal and a more vertical ventral area. http://purl.obolibrary.org/obo/HAO_0001886 frontal ledge
head The tagma that is located anterior to the thorax. http://purl.obolibrary.org/obo/HAO_0000397 head
hyperoccipital carina The carina that extends on the vertex between the outer orbits. http://purl.obolibrary.org/obo/HAO_0000406 hyperoccipital carina
integument The anatomical system that forms the covering layer of the animal, ectodermal in origin and composed of epidermal cells producing the cuticle. http://purl.obolibrary.org/obo/HAO_0000421 integument
lateral ocellus The ocellus that is paired. http://purl.obolibrary.org/obo/HAO_0000481 lateral ocellus
lateral propodeal carina The carina that arises submedially from the anterior margin of the metapectal-propodeal complex, is longitudinal and extends towards the propodeal foramen. http://purl.obolibrary.org/obo/HAO_0001919 lateral propodeal carina
malar striae The anatomical cluster posterior to the malar sulcus that is composed of carinae radiating from the pleurostomal condyle. http://purl.obolibrary.org/obo/HAO_0002373 malar striae
malar sulcus The sulcus that extends between the ventral margin of the compound eye and the base of the mandible. http://purl.obolibrary.org/obo/HAO_0000504 malar sulcus
mandible The appendage that is encircled by one sclerite that is connected to the cranium proximolaterally and to the maxillo-labial complex proximomedially via conjunctivae and articulates with the cranium via the anterior and posterior cranio-mandibular articulations. http://purl.obolibrary.org/obo/HAO_0000506 mandible
margin The line that delimits the periphery of an area. http://purl.obolibrary.org/obo/HAO_0000510 margin
mesepimeral ridge The ridge that extends along the posterior margin of the mesopectus. http://purl.obolibrary.org/obo/HAO_0000537 mesepimeral ridge
mesepimeral sulcus The sulcus that extends along the posterior margin of the mesopectus, delimits the mesepimeral area and corresponds to the mesepimeral ridge. http://purl.obolibrary.org/obo/HAO_0000538 mesepimeral sulcus
mesopleural carina The carina that crosses the mesopleuron and limits ventrally the femoral depression. http://purl.obolibrary.org/obo/HAO_0000559 mesopleural carina
mesopleural epicoxal sulcus The epicoxal sulcus that is located on the mesopleuron. http://purl.obolibrary.org/obo/HAO_0000560 mesopleural epicoxal sulcus
mesopleural pit The pleural pit that is located on the mesopleuron. http://purl.obolibrary.org/obo/HAO_0000561 mesopleural pit
mesopleuron The pleuron that is located in the mesothorax. http://purl.obolibrary.org/obo/HAO_0001354 mesopleuron
mesoscutal humeral sulcus The sulcus that extends medially along the parascutal carina and corresponds to a shallow ridge. http://purl.obolibrary.org/obo/HAO_0000569 mesoscutal humeral sulcus
mesoscutal suprahumeral sulcus The sulcus that extends along the anterior margin of the mesoscutum between the anterior-most point of the preaxilla and the anteroadmedian line and corresponds to the vertical lobe of the mesoscutum. http://purl.obolibrary.org/obo/HAO_0000570 mesoscutal suprahumeral sulcus
mesoscutum The scutum that is located on the mesonotum. http://purl.obolibrary.org/obo/HAO_0000575 mesoscutum
mesosoma The anatomical cluster that is composed of the prothorax, mesothorax and the metapectal-propodeal complex. http://purl.obolibrary.org/obo/HAO_0000576 mesosoma
metanotal trough The area that is concave, and is delimited medially by the metascutellum, laterally by the supraalar area and posteriorly by the metascutellar arm. http://purl.obolibrary.org/obo/HAO_0000600 metanotal trough
metapleural carina The carina that delimits the metapleuron dorsally from the propodeum, extends from just ventral of the metapleural arm to the metacoxal articulation and passes anteroventral to the propodeal spiracle. http://purl.obolibrary.org/obo/HAO_0000609 metapleural carina
metapleural sulcus The line that corresponds with the metapleural ridge. http://purl.obolibrary.org/obo/HAO_0000614 metapleural sulcus
metapleuron The area of the metapectal-propodeal complex that is located laterally of the metadiscrimen. http://purl.obolibrary.org/obo/HAO_0000621 metapleuron
metascutellum The area that is located posteromedially on the metanotum, is delimited laterally by the metanotal trough and corresponds to the reservoir of the dorsal vessel. http://purl.obolibrary.org/obo/HAO_0000625 metascutellum
metasoma The tagma that is connected anteriorly to the metapectal-propodeal complex at the propodeal foramen and consists of abdominal segments. http://purl.obolibrary.org/obo/HAO_0000626 metasoma
metasomal depression The acetabulum that is concave, surrounds the nucha and accommodates the base of the metasoma. http://purl.obolibrary.org/obo/HAO_0000627 metasomal depression
metasomal segment The abdominal segment that is located in the metasoma. http://purl.obolibrary.org/obo/HAO_0001969 metasomal segment
netrion The area that is located posteroventrally on the pronotum and corresponds to the site of origin of first flexor of the fore wing muscle. http://purl.obolibrary.org/obo/HAO_0000644 netrion
netrion sulcus The sulcus that anteriorly delimits the netrion. http://purl.obolibrary.org/obo/HAO_0000646 netrion sulcus
notaulus The line that extends submedially along the mesoscutum and corresponds to the median border of the site of origin of the first mesopleuro-mesonotal muscle. http://purl.obolibrary.org/obo/HAO_0000647 notaulus
occipital carina The carina that surrounds dorsolaterally the occiput. http://purl.obolibrary.org/obo/HAO_0000653 occipital carina
ocellar diameter The diameter of the ocellus. http://purl.obolibrary.org/obo/HAO_0002107 ocellar diameter
ocellus The multi-tissue structure that is located on the top of the head, composed of the corneal lens, pigment cell, rhabdoms and synaptic plexus. http://purl.obolibrary.org/obo/HAO_0000661 ocellus
orbital carina The carina that is located on the face, parallels the inner orbit and is paired. http://purl.obolibrary.org/obo/HAO_0000810 preorbital carina
ovipositor The anatomical cluster that is composed of the first valvulae, second valvulae, third valvulae, first valvifers, second valvifers and female T9. http://purl.obolibrary.org/obo/HAO_0000679 ovipositor
postacetabular sulcus The sulcus that extends posteriorly along the epicnemial carina. http://purl.obolibrary.org/obo/HAO_0000741 postacetabular sulcus
posterior mesoscutellar sulcus The line that extends along the posterior margin of the mesoscutellum and corresponds to the posterior mesoscutellar ridge. http://purl.obolibrary.org/obo/HAO_0000757 posterior mesoscutellar sulcus
posterior pronotal inflection The inflection that extends along the posterior margin of the pronotum and articulates with the anterior mesopleural inflection. http://purl.obolibrary.org/obo/HAO_0000761 posterior pronotal inflection
postmarginal vein The abscissa that is marginal and located distal to the marginal vein. http://purl.obolibrary.org/obo/HAO_0000783 postmarginal vein
prespecular sulcus The sulcus that delimits anteriorly the speculum and corresponds to the anterior margin of the speculum. http://purl.obolibrary.org/obo/HAO_0000816 prespecular sulcus
pronotal cervical sulcus The sulcus that extends along the anterior margin of the pronotum and delimits the anterior rim of pronotum. http://purl.obolibrary.org/obo/HAO_0000831 pronotal cervical sulcus
pronotum The notum that is located in the prothorax. http://purl.obolibrary.org/obo/HAO_0000853 pronotum
propleural epicoxal sulcus The epicoxal sulcus that sets of the epicoxal lobe from the ventral part of the ventral propleural area. http://purl.obolibrary.org/obo/HAO_0000858 propleural epicoxal sulcus
propleuron The pleuron that is articulated with the fore leg, connected dorsolaterally (anterolaterally) with the pronotum and ventrally (posteriorly) with the prosternum. http://purl.obolibrary.org/obo/HAO_0000862 propleuron
propodeal carina The carina that is located on the propodeum. http://purl.obolibrary.org/obo/HAO_0000864 propodeal carina
propodeum The area of the metapectal-propodeal complex that is located posterior to the metapleural carina. http://purl.obolibrary.org/obo/HAO_0001249 propodeum
sculpture The area that is located on the sclerite and that is composed of repetitive anatomical structures. http://purl.obolibrary.org/obo/HAO_0000913 sculpture
scutoscutellar sulcus The sulcus that extends along the scutoscutellar suture. http://purl.obolibrary.org/obo/HAO_0000919 scutoscutellar sulcus
segment An anatomical structure that is metameric and is connected to other metameric subdivisions by muscles and is delimited by its sclerites. http://purl.obolibrary.org/obo/HAO_0000929 segment
sensillum A sense organ embedded in the integument and consisting of one or a cluster of sensory neurons and associated sensory structures, support cells and glial cells forming a single organized unit with a largely bona fide boundary. http://purl.obolibrary.org/obo/HAO_0000933 sensillum
skaphion The area that is anteriorly on the mesonotum and delimited posteriorly by the skaphion carina. http://purl.obolibrary.org/obo/HAO_0000940 skaphion
spiracle The anatomical cluster that is composed of the distal end of the trachea and the margin of the sclerite or conjunctiva surrounding the spiracular opening. http://purl.obolibrary.org/obo/HAO_0000950 spiracle
sternite The sclerite that is located on the sternum. http://purl.obolibrary.org/obo/HAO_0000955 sternite
stigmal vein The vein that is adjacent proximally to the pterostigma. http://purl.obolibrary.org/obo/HAO_0002428 stigmal vein
submarginal vein Basal-most portion of the forewing vein complex that occurs behind the costal cell; measured from the constriction that delimits the humeral plate to the point at which the vein touches the leading edge of the wing apically. http://purl.obolibrary.org/obo/HAO_0000972 submarginal vein
sulcus The groove that corresponds to a ridge. http://purl.obolibrary.org/obo/HAO_0000978 sulcus
tergite The sclerite that is located on the tergum. http://purl.obolibrary.org/obo/HAO_0001005 tergite
trachea The cuticular invagination that is tubular, branched into tracheoles and bears the taenidia. http://purl.obolibrary.org/obo/HAO_0002415 trachea
transepisternal line The line that is longitudinal, extends ventrolaterally on the mesopleuron and corresponds with the site of origin of the second and third mesopleuro-third axillary sclerite of fore wing muscle and the second mesopleuro-mesonotal muscle. http://purl.obolibrary.org/obo/HAO_0001205 transepisternal line
transverse pronotal carina The carina that delimits posteriorly the pronotal neck. http://purl.obolibrary.org/obo/HAO_0001031 transverse pronotal carina
venation The anatomical cluster that is composed of abscissae. http://purl.obolibrary.org/obo/HAO_0001096 venation
ventral metapleural area The area that is located on the metapleuron anteroventrally of the metapleural sulcus. http://purl.obolibrary.org/obo/HAO_0001062 ventral metapleural area
wing The appendage that is between the notum and the pectus and is connected to the body by the axillary sclerite muscles. http://purl.obolibrary.org/obo/HAO_0001089 wing