Systematics of Trichoteleia Kieffer and Paridris Kieffer (Hymenoptera, Platygastroidea, Platygastridae)

Paridris Kieffer and Trichoteleia Kieffer are morphologically similar genera of solitary egg parasitoids with little overlap between their distributions: Paridris is found commonly worldwide with the exceptions of Madagascar, from which a single specimen is known, and New Zealand, from which no records are known; Trichoteleia is endemic to the Malagasy islands. Here we present the first phylogenetic analysis of platygastroid wasps that combines and compares morphological and molecular data. We find the results of the phylogenetic analyses of the two data sources to be largely congruent for the species treated here. Paridris and Trichoteleia are found to be monophyletic, as are two morphologically well-defined species groups within Paridris. Neoparidris Galloway is found to belong within Paridris and is treated as a junior synonym, syn. n. The faunas of Paridris from Africa, Melanesia and the Indo-Malay islands are revised. Fifteen species are described of which 9 are new: Paridris anikulapo Talamas, sp. n. (sub-Saharan Africa); Paridris densiclava (Kieffer), (Seychelles); Paridris bispinosa (Masner), (Gabon); Paridris nigriclava (KiefJHR 34: 1–79 (2013) doi: 10.3897/JHR.33.4714 www.pensoft.net/journals/jhr Copyright Elijah J. Talamas et al. This is an open access article distributed under the terms of the Creative Commons Attribution License 3.0 (CC-BY), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. MonogrAPH Elijah J. Talamas et al. / Journal of Hymenoptera Research 34: 1–79 (2013) 2 fer), (Seychelles); Paridris nitidiceps (Kieffer), (Seychelles); Paridris tenuis (Nixon), (sub-Saharan Africa); Paridris trispinosa Talamas & Masner, sp. n., (Cameroon, Democratic Republic of the Congo); Paridris bifurcata (Dodd), comb. n., (Australia, Papua New Guinea); Paridris mnestros Talamas & Masner, sp. n., (Indonesia, Malaysia); Paridris pantex Talamas, sp. n., (Fiji); Paridris phrikos Talamas & Masner, sp. n., (Fiji); Paridris skolops Talamas & Masner, sp. n., (Fiji); Paridris sulcata Talamas, sp. n., (Vanuatu); Paridris taekuli Talamas & Masner, sp. n., (Australia, Bangladesh, Fiji, India, Indonesia, Ivory Coast, Madagascar, New Caledonia, Thailand, Vietnam); Paridris xestos Talamas & Masner, sp. n., (Fiji). Paridris flaviclava (Kieffer), syn. n., and P. nigraticeps (Kieffer), syn. n., are treated as junior synonyms of P. nigriclava.


Introduction
Paridris is a genus of minute wasps that, extrapolating from a single host record from North America (Masner and Muesebeck 1968), are parasitoids of cricket eggs (Orthoptera: Gryllidae). The genus is nearly cosmopolitan in distribution: it is not known from New Zealand and its presence in Madagascar is known from a single specimen. Significantly, the putative sister group to Paridris, Trichoteleia, is endemic to the Malagasy Islands. The nearly exclusive distributions of these genera and their morphological similarity suggested the possibility that Trichoteleia was an apomorphic lineage derived from within Paridris. To test this hypothesis we conducted a phylogenetic analysis of these genera based on molecular and morphological data. This is the culmination of our examination of these groups of parasitoids, following our revision of Trichoteleia at the species level and evaluation of its generic limits (Talamas et al. 2011a), a generic level assessment of Paridris, and species descriptions of the nephta group (formerly the genus Tuora, Talamas et al. 2011b) and the New World species of Paridris (Talamas et al. 2012).
Among platygastroid genera, the diversity of species revealed by recent taxonomy is often an order of magnitude greater than was previously known (e.g., Johnson et al. 2008, Taekul et al. 2008, and Paridris is no exception. Revision of the Paridris nephta species group increased the number of species from 1 to 15 (Talamas et al. 2011b), and treatment of the New World fauna resulted in a similar increase from 2 valid species to 15 (Talamas et al. 2012). Here we continue our revision of Paridris with two goals. First, we strive to examine the gamut of morphological diversity within the genus to produce a maximally informed coding scheme for phylogenetic characters and form an accurate, robust generic concept. Second, we seek to document the species level diversity and distribution of Paridris and produce identification tools that make these data usable for future biological studies. We present the following as a single publication because we consider it best to make taxonomic decisions, such as the synonymy of Neoparidris, in a phylogenetic context.
Our focus for this revision is on the geographical regions of Africa, Melanesia and the Indo-Malay islands based on the accessibility of primary types. Kononova and Kozlov (2008) produced a key to the species of the Palearctic, Rajmohana (2007) published a key to the species of India, and Kozlov and Lê (2000) published a key to the species of Vietnam. Together with the present work, these publications treat most of the world's geographic areas. Only the fauna of Australia remains largely unexplored.
Although we were unable to access the type material for the Indian species, the high quality images of Paridris spinosa Rajmohana and the key to Indian species (Rajmohana 2007) allowed us to identify this species and P. armigera among the material at hand and include them in our analysis. Our analyses include a species from Southeast Asia that was likely described by Kozlov and Lê (2000). We were unable to unambiguously identify this species with their key and it is indicated as Paridris asian sp. 1. Synopses of the species not analyzed taxonomically or phylogenetically by the present authors may be found following the species descriptions.
Previous phylogenetic analyses of platygastroids have used morphological data (Iqbal andAustin 2000, Valerio et al. 2010) or molecular data (Carey et al. 2006, Murphy et al. 2007), but to date none have compared the two datatypes with the same set of taxa or conducted a combined analysis. Here we demonstrate the utility of morphological characters at the species level, and to some extent at the generic level. We show that morphological characters, though demonstrably homoplasious at times, are useful for reconstructing relationships, particularly when combined with molecular data. These analyses represent the first comparison between analyses of molecular and morphological data within the superfamily.
This work is conducted as part of the Platygastroidea Planetary Biodiversity Inventory and represents a step toward a species-level revision of the Scelionini sensu lato. The contributions of the authors are as follows: E.J. Talamas: DNA extraction and amplification; sequence alignment and phylogenetic analysis, character definition and coding, species concept development, imaging, key development, manuscript preparation; L. Masner: aggregation of specimens, species concept development, manuscript preparation; N.F. Johnson: software and database development, character definition; manuscript preparation.

Materials and methods
Primary types: The primary types of J. J. Kieffer and G. E. J. Nixon in The Natural History Museum were photographed by E. Talamas during a visit to this collection in 2009. Our assessment of Neoparidris, and ultimately its treatment as a junior synonym of Paridris, was facilitated by images of the type species taken by N. F. Johnson in 2004 at the Queensland Museum, Brisbane, Australia. Continual access to images of the type material made this project possible without risking damage to specimens during shipping. We hope that this demonstration of the utility of such photographs will encourage the imaging of type material as standard practice in taxonomy.
Morphological terms used in this revision were matched to the Hymenoptera Anatomy Ontology (HAO, Yoder et al. 2010) (Appendix I). Identifiers (URIs) in the format http://purl.obolibrary.org/obo/HAO_XXXXXXX represent anatomical concepts in HAO version http://purl.obolibrary.org/obo/hao/2011-05-18/hao.owl. They are provided to enable readers to confirm their understanding of the anatomical structures being referenced. To find out more about a given structure, including, images, references, and other metadata, use the identifier as a web-link, or use the HAO:XXXXXXX (note colon replaces underscore) as a search term at http://glossary.hymao.org.
The description of surface sculpture is presented in two formats. Areas of the exoskeleton in which the sculptural elements are inseparable are described simply as "sculpture". For areas in which the sculptural elements vary independently, sculpture is divided into three categories: punctation: round depressions associated with setae; macrosculpture: raised or sunken patterns of texture that are oriented linearly or radially with respect to punctation or the axes of the body; microsculpture: unoriented, very fine wrinkles or pustulations that occur on, in, or between elements of macrosculpture and punctation.
Information management: The locality data reported for primary types are not a literal transcription of the labels: some abbreviations are expanded; additional data from the collectors are also included. The holotypes should be unambiguously identifiable by means of the unique identifier or the red holotype label. The numbers prefixed with "OSUC " and "CASENT " are unique identifiers for the individual specimens (note the blank space after the 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. This monograph also features simultaneous publication and distribution of taxonomic and occurrence records through the Global Biodiversity Information Facility (GBIF) using DarwinCore Archives. All new species have been prospectively registered with Zoobank (Polaszek et al. 2005) and other taxonomic names have been retrospectively registered therein. All names are also registered in the Hymenoptera Name Server (hns.osu.edu). Life sciences identifiers, lsids, may be resolved at the URLs specified in the footnotes or at lsid.tdwg.org.
Cybertools: The species descriptions are generated by a database application, vS-ysLab (purl.oclc.org/NET/hymenoptera/vSysLab), designed to facilitate the generation of taxon by character data matrices, to integrate these with the existing taxonomic and specimen-level database, and to export the data both as text and as input files for other applications (Johnson 2010). The output is in the format of "Character: Character state(s)." Intraspecific variability is indicated by character states separated by a semicolon. The illustrated matrix of morphological characters used in our phylogenetic analysis can be found at http://vsyslab.osu.edu/show_matrix.html?project_id=106.
Imaging: Images were produced using Combine ZP and AutoMontage extendedfocus software. The individual images are archived at the image database at The Ohio State University (purl.oclc.org/NET/hymenoptera/specimage) and with MorphBank (www.morphbank.net). The latter also contains collections of images organized by plate.
Species concept: For the purpose of this revision, species are defined as taxa diagnosable by putative autapomorphies or a unique combination of fixed character states.
Molecular data: DNA was extracted nondestructively with a Qiagen DNeasy extraction kit and amplified according to standard protocols with the primers of Murphy et al. (2007). Sequences of ribosomal genes were aligned by eye according to the structural models of  and . CO1 sequences, and the variable loop regions of 18S and 28S, were aligned with MUSCLE (Edgar 2004). The CUIDs of voucher species and Genbank accession numbers are presented in Appendix IV.
Phylogenetic analysis: We analyzed our data under the criterion of parsimony using TNT (Goloboff et al. 2008) with gaps treated as missing data in all analyses and equal weights for all characters. We consider parsimony to be the optimal method for our dataset because it contains both morphological and molecular data, and the latter are missing for two thirds of the species, creating a pitfall for parameter estimation in model-based analyses. Parsimony also enables comparison between the signal in morphological and molecular data within the same analysis paradigm. We used the script of Peña et al. (2006) to perform a Partitioned Bremer Support analysis in TNT with four data partitions, morphology, 28S, 18S and CO1, to examine the contributions of each data set to support for the nodes in the strict consensus tree. The composite consistency and retention indices (CI and RI) are listed in the figure captions for each phylogeny. The matrix used for phylogenetic analysis is included as Appendix V.
Composite terminals: The CO1 sequence for our outgroup terminal, Archaeoteleia, was amplified from Archaeoteleia mellea, and 18S and 28S sequences from A. onamata. Morphology was coded from A. mellea. For the morphological characters used in our analysis, these species of Archaeoteleia are essentially isomorphic. The sequence data for Paridris aeneus came from two specimens: OSUC 261872 for CO1 and OSUC 265221 for 18S and 28S.
Excluded species: We excluded P. armata Talamas, P. invicta Talamas, P. nitidiceps and P. densiclava from our final analyses. Paridris armata, P. invicta, and P. nitidiceps are known only from males, lacking phylogenetically important female characters, and are of uncertain affinity. The morphological characters of P. nitidiceps and P. densiclava were coded from photographs of the type specimens. Consequently, we were unable to observe a number of the characters that we consider to be phylogenetically informative. Apart from a loss of resolution, analyses that included these species did not differ from those presented in Figs 2-3.

results of phylogenetic analysis
All of our analyses confirmed monophyly of the P. nephta and P. pallipes species groups and Trichoteleia (Figs 1-3) with strong bootstrap support for these clades in the molecular and combined analyses. Trichoteleia and the P. nephta group had significant support in the morphological analysis, but support for the P. pallipes group here was poor, reflecting the homoplasious nature of the characters that delimit this group. Similarly, Paridris was retrieved as a monophyletic group in all of the analyses, but its highest bootstrap value of 49, retrieved in the analysis of combined data, is still low. From a morphological perspective this is unsurprising because all of the synapomorphies for Paridris are also found in other genera or are lost secondarily. The presence of Probaryconus sp. 2 among Paridris in the morphological analysis ( Fig. 2) is also not unexpected, particularly because this specimen was selected for its similarity to Paridris: it lacks an epomial carina and has setose compound eyes. Topologically, the only character that separates this species from Paridris is the externally undifferentiated metascutellum. Even this character must be carefully assessed because in some species of both Paridris and Probaryconus the horn of T1 may be very large and preclude observation of the metanotum. However, the high bootstrap support for Probaryconus in the molecular and combined analyses ultimately affirms confidence in our concepts for these genera. A subset of characters that are diagnostic for genera and species treated in this analysis (see Appendix II) are mapped onto the combined data phylogeny in Figure 3.
The analysis of Partitioned Bremer Support (see Appendix III) indicates that contributions to clade support from the four data partitions, morphology, 28S, 18S, and CO1, are clade dependent. Within Trichoteleia, 28S provided minimal clade support with values of zero for most of the nodes. CO1 contained the most contrarian signal for this genus and accounted for nearly all of the disagreement between partitions with nega-tive values for six of the twenty nodes. However, none of these were less than negative one, indicating that the incongruence of CO1 with the other data partition is small in magnitude for Trichoteleia. The P. nephta species group yielded a similar pattern with no contribution to node support from 28S and a small degree of incongruence from CO1 and 18S. At each node within this group morphology provided the strongest signal.
The pattern of clade support from 18S was consistent with a relatively slow rate of evolution in this gene; nodes at the base of Paridris had 18S support values an order of magnitude higher than those toward the tips. Within the the Scelio+Calliscelio clade the support values for all three genes were the largest (both positive and negative) with the signal of 18S and CO1 conflicting with, and overriding, that of 28S.

Discussion
Morphological homoplasy within Platygastroidea has been mentioned by previous authors Huggert 1989, Iqbal andAustin 2000) and it is present in our data as well. However, Paridris, for which no uncontroverted synapomorphy exists in our data set, nonetheless formed a clade in all of our analyses, with the caveat that Probaryconus sp. 2 is present in this clade in the analysis of morphology alone. Our reductionist coding system (vs. composite characters) may eliminate some important characters that contribute to this erroneous placement. Specifically, the pattern of punctation and microsculpture on the mesoscutum of Probaryconus is found throughout this genus and is recognizable to the experienced taxonomist, but when broken into component characters the characters no longer define Probaryconus as a group because of the dif-   ficulty in accurately coding and articulating subtly different forms of microsculpture. We emphasize that the use of morphological data can be extremely useful by allowing the inclusion of taxa for which molecular data is not available.
We note that the genera within Scelioninae are typically well-defined groups, but are based on unique combinations of characters found throughout the subfamily, that are thus homoplasious in phylogenetic analyses. In polytypic groups, such as Paridris, even the characters that define the group are either secondarily lost (transverse carina on T2), or are apparently homoplasious (metascutellum). Trichoteleia is a group well defined by multiple synapomorphies (felt fields of T1 and S2, setose metascutellum), but these characters do little to ally it with other genera. For example, setation of the eyes is common, and may be present or absent within genera (Idris, Probaryconus). Metascutellar setation, though uncommon, is found in P. spinosa and P. taekuli as well as in genera that are morphologically more distant (Chromoteleia, Bracalba, Sceliacanthella (OSUC 150176)) and in a species of Teleasinae (OSUC 281605).
The outgroups in our analysis represent a small fraction of the subfamily Scelioninae, and we do not conclude from these results any relationships at the level of genera, only that Trichoteleia is not derived from within Paridris, answering our primary question.

Paridris pallipes species group
The P. pallipes species group is morphologically distinct from the remainder of Paridris, noticeable immediately by the relative absence of macrosculpture from the head and mesosoma. The geographical distribution of this group is perplexing-it is found throughout North and South America and in the Fijian Islands. This suggested the possibility of "tramp" species, yet no species are shared between the two regions. Additionally, P. pantex (Fiji) has a highly apomorphic form of the felt fields on S2 that we consider unlikely to have evolved during recent history in which humans have been able to travel rapidly between Fiji and the Americas. It is possible that the group was once widespread, and the distribution we see now is the result of extinction, or that one of the centers of diversity is simply a radiation of the other. Either way, our understanding of the group, and Paridris as a whole, will be greatly furthered by additional host and biological data that allows us to make more informed inferences.
The association between the P. pallipes species group and islands is noteworthy. In addition to the three species known from the Fijian islands, 6 of the 8 species of the P. pallipes group in the New World are found on Caribbean islands, 4 of them exclusively so.
Diagnosis. The P. pallipes species group can be separated from the remainder of Paridris by the following combination of characters: genal striae strongly reduced, rarely extending to midpoint of compound eye; occipital carina absent below foramen magnum; occipital carina complete dorsally; dorsal frons and vertex without macrosculpture; plical carina absent; posterior margin of metascutellum straight to convex; antecostal sulcus of T2 present as a constriction or line of foveae, without carina along its posterior margin; postmarginal vein punctiform.
In addition to these ubiquitously present characters, species of the P. pallipes group often have dense setation on the postgena and S1. All species except for P. pantex have the felt field present as a line of dense setae along a longitudinal ridge. In a few specimens of P. dnophos and P. pallipes the lateral ocellus is less than two ocellar diameters from the inner orbit of the compound eye. However, in the vast majority of specimens of these species, and in all other members of this species group, the lateral ocellus is distinctly remote from the inner orbits.

African Paridris
The fauna of Paridris in continental Africa is surprisingly small with just five species. Two of these, P. tenuis and P. anikulapo, are widespread in distribution and found in eastern, western and southern Africa. Our knowledge about their presence in central Africa, and the existence of other species of Paridris, is currently limited by a dearth of collecting in this region.
Three valid species are known from the Seychelles. Paridris densiclava and P. nitidiceps were described by J. J. Kieffer from singletons of opposite sex. We have no additional material of either species, and because we found characters to separate them we consider it best to keep them as separate species. However, we acknowledge that we are currently unable to assess intraspecific variation, and that examination of more material may reveal them to be conspecific.
The single species from Madagascar, P. taekuli, is known from the Ivory Coast, South and Southeast Asia, Fiji, New Caledonia and Northern Australia from a modest number of specimens. Its sister species in the New World, P. psydrax, ranges from Argentina to California and is similarly known from a rather short series given its wide distribution. Gena along posterodorsal margin of eye smooth and shining ( Fig 28); notaulus percurrent (Fig. 29)  Occipital carina incomplete and not reaching base of mandible (Fig 8-9); postmarginal vein less than half as long stigmal vein (Fig. 14)  Color of head: brown to black. Distal margin of clypeus: serrate. Shape of distal margin of clypeus in anterior view: straight. Width of clypeus: greater than width across toruli. Lateral corner of clypeus: projecting into acute angle. Length of mediofacial striae: not extending above midpoint of compound eye. Anterodorsal node on interantennal process: absent. Central keel: absent. Length of OOL: less than 2 ocellar diameters. Macrosculpture of frons between median ocellus and inner orbit of eye: dorsoventrally strigose; absent. Patch of microsculpture posterior to lateral ocellus in male: absent. Diagnosis. Paridris anikulapo is closest to P. bispinosa, and may be separated by the smooth and shining gena along the posterodorsal orbit of the compound eye.

Key to African
Etymology. The species epithet "anikulapo" is a Yoruban name meaning "he who carries death in his pouch". It is given to this species as a reference to the parasitoid life history and is treated as a noun in apposition.
Link Number of basiconic sensilla on A8: one. Shape of male flagellomeres: longer than wide by a factor less than 2.
Color of head: black; reddish brown. Distal margin of clypeus: serrate. Shape of distal margin of clypeus in anterior view: convex. Width of clypeus: greater than width across toruli. Lateral corner of clypeus: projecting into acute angle. Length of mediofacial striae: continuous with sculpture of dorsal frons. Anterodorsal node on interantennal process: present. Central keel: absent. Length of OOL: less than 2 ocellar diameters. Macrosculpture of frons between median ocellus and inner orbit of eye: Diagnosis. In P. bispinosa, the metascutellum is distinctly bispinose and T6 is sharply constricted in its apical half, separating it from all but two African species, P. anikulapo and P. trispinosa. The females of P. bispinosa have the notaulus present as a single fovea on the posterior margin of the mesoscutum and in females of P. anikulapo the notaulus extends to the anterior mesoscutum. In the specimen of P. bispinosa examined here, the gena is coarsely sculptured throughout and in P. anikulapo the gena along the posterior margin of the eye is smooth and shining. Paridris trispinosa may be separated from P. bispinosa by the presence of a posteriorly directed spine on the horn of T1.

Comments.
We did not examine any males of P. bispinosa in this revision, but we speculate that they will have an abbreviate notaulus, as in the females of this species, and that this will enable separation from males of P. trispinosa. ( Anterior projection of the propodeum: absent. Setation of metasomal depression: absent. Posterior projection of the propodeum: present as a point formed by plical and lateral propodeal carinae. Plical carina: present. Lateral propodeal area: raised above propodeal surface and indicated by lesser setation. Shape of lateral propodeal area: con-tinuous with prespiracular propodeal area. Sculpture of lateral propodeal area: weakly to moderately rugose.

Paridris densiclava
Length of postmarginal vein: equal to stigmalis. Rs in fore wing: spectral. Cu vein in fore wing: spectral. M vein in forewing: spectral. Color of costal cell in female: hyaline. Color of sub-radial area in female: hyaline. Color of cubito-medial area in female: hyaline. Color of anal margin in female: hyaline. RS+M in forewing: nebulous. Basal vein in hind wing: spectral. Setation of hind wing: uniform throughout.
Color of metasoma: reddish brown. Longitudinal median carina on horn of T1: absent. Armature on posterior surface of T1 horn: absent. Patch of dense fine setae on anterolateral T1: absent. Constriction of apical T6 in female: absent.
Diagnosis. Paridris densiclava shares the smoothly convex shape of T6 with P. nigriclava, and differs by having a postmarginal vein as long as the stigmal vein. This venation is shared by P. nitidiceps, also from the Seychelles and known from a single male. We separate these species on the basis of the complete notaulus and punctate posterior mesepimeral area in P. nitidiceps. The notaulus of P. densiclava is present as a single fovea on the posterior margin of the mesoscutum and the posterior mesepimeral area is entirely smooth.
Number of basiconic sensilla on A8: two. Shape of male flagellomeres: longer than wide by a factor less than 2.
Color of head: brown to black; reddish brown. Distal margin of clypeus: serrate. Comments. The posterior margin of the metascutellum, a character used previously in identification keys for this species, is typically emarginate, but may be straight or convex. Talamas  Number of basiconic sensilla on A8: two. Shape of male flagellomeres: longer than wide by a factor less than 2.

Paridris trispinosa
Color of head: dark brown. Distal margin of clypeus: serrate. Shape of distal margin of clypeus in anterior view: convex. Width of clypeus: greater than width across toruli. Lateral corner of clypeus: projecting into acute angle. Length of mediofacial striae: not extending above midpoint of compound eye; continuous with sculpture of dorsal frons. Anterodorsal node on interantennal process: absent. Central keel: absent. Length of OOL: less than 2 ocellar diameters. Macrosculpture of frons between median ocellus Diagnosis. Paridris trispinosa is most similar to P. bispinosa. Females of P. trispinosa may be separated by the posteriorly directed spine on the horn of T1 (absent in P. bispinosa) and percurrent notauli (abbreviate in P. bispinosa). Among the specimens examined in this revision, the surface sculpture of P. trispinosa is coarser than that of P. bispinosa, particularly on the lateral mesoscutum.
Etymology. The Latin adjectival epithet "trispinosa" is given to this species for the spines of the metascutellum and horn of T1. Comments. We did not examine any males of P. bispinosa in this revision, but we speculate that they will have an abbreviate notaulus, as in the females of this species, and that this will enable separation from males of P. trispinosa.

Paridris of Melanesia and the Indo-Malay Islands
From the islands of Borneo and Sulawesi we describe a single species. Although this region has few species of Paridris, it harbors a species, P. mnestros, which is important for understanding morphological diversity in this genus. Its form of the felt field on S2 is unique, and it is the only species outside the P. nephta group with bright and patterned coloration of the body.
With five species, the Fijian islands are a hotspot of diversity for Paridris. Three of these species belong to the P. pallipes species group, a lineage otherwise known only from the New World.
In addition to the specimens of P. bifurcata, we examined a single male specimen (OSUC 265189) from Papua New Guinea that exhibits characteristics of the P. pallipes species group and does not belong to any of the species treated here. We believe that mention of this species is noteworthy, but we choose not to describe it on the basis of single male specimen outside the context of a comprehensive revision of the species from mainland Southeast Asia and Australia. Horn of T1 with posteriorly directed spine (Fig. 80); A8 with 1 basiconic sensillum (as in Fig. 5)  Felt field of S2 present anteromedially (Figs 18-19); notaulus straight posteriorly (Fig. 11, 67); anterior propodeal projection absent or indicated by weak protuberance (Fig. 13) (Figs 20-21); notaulus expanded posteriorly (Fig. 10); anterior propodeal projection present as conspicuous point or spine (Figs 12, 75)  Horn of T1 smooth or with median longitudinal carina (Fig. 90, 92); T3 without surface sculpture (Fig. 95) (Fig. 75); T3 longitudinally striate laterally (Fig. 67) Posterior margin of metascutellum straight or convex (Figs 12-13, 79); occipital carina absent or not extending below foramen magnum (Figs 8-9) ... 3 2 S2 felt field present anterolaterally in coarsely rugose excavation (Fig. 21); metasoma banded (Fig. 62) ... Paridris mnestros Talamas & Masner, sp. n. -S2 felt field present laterally as a longitudinal patch of setigerous punctures (Fig. 23); metasoma black (Fig. 61)  Felt field of S2 present anteromedially (Figs 18-19); notaulus straight (Figs 11,67); anterior propodeal projection absent or indicated by weak protuberance (Fig. 13) (Fig. 10); anterior propodeal projection present as conspicuous point or spine (Fig. 12)   Comments. Galloway (1984) erected Neoparidris to accomodate a species that had attributes of Paridris but lacked the small eyes, compact ocellar triangle, and incomplete notaulus that characterized his concept of the group. In his discussion of Neoparidris, Galloway commented that this species may ultimately belong in Paridris when its limits became clearer, and this is indeed the case. Talamas  Diagnosis. Paridris mnestros uniquely has the felt field of S2 as an anteriorly located pit of coarse rugae. The constriction of the apex of T6 into a rather sharp point is also not found in any other species of Paridris. Lastly, this is the only species outside of the P. nephta species group to exhibit bright coloration of the body and a banded metasoma.

Etymology. The
Greek word for marriage "mnestros" is given to this species to commemorate the wedding of Ryan St. Clair to Grace Wong, and because this species combines morphological structures commonly found in Paridris (i.e. apical constriction of T6, posteriorly directed spine on horn of T1) with the coloration, mesoscutal surface sculpture, and very long setation found in the P. nephta species group.
Link to distribution map. Diagnosis. Paridris pantex may be distinguished from all known species in this genus by the dense tufts of setae on anteromedial S2.
Etymology. The epithet "pantex", meaning "paunch" is given to this species for the dense tufts of setae on anteromedial S2. The name is treated as a noun in apposition.
Link to distribution map. 33 Material examined. Holotype, female: FIJI: Northern Div., Cakaudrove Prov., Taveuni Isl., 5.6km SE Tavuki, MT1, Devo Peak, 16.843°S 179.966°W, 1187m, 14.XI-21.XI.2002  Diagnosis. Paridris phrikos is most similar to P. xestos and females may be separated by the presence of a transverse ridge on the horn of T1 and longitudinal striation on lateral T3. Males lack this character and are separated from those of P. xestos solely on the basis of having weak striation on lateral T3. For this reason, males are excluded from the paratype series.
Etymology. The Greek word "phrikos", meaning "ruffling of a smooth surface", refers to the transverse ridge on the horn of T1. The name is treated as a noun in apposition.
Link to distribution map. Diagnosis. Paridris skolops is unique among the Paridris species in Fiji because the female has a carina along the posterior margin of the antecostal sulcus on T2. Males and females have a setose metasomal depression and dense fine setae along the postgena which serve to separate them from the other species treated in this revision.
Etymology. The Greek epithet "skolops" meaning "anything pointed" refers to the posteriorly directed spine on the horn of T1 in females of this species. The name is treated as a noun in apposition.
Link Diagnosis. P. sulcata is morphologically closest to P. bifurcata. Females may be separated by the posteriorly directed spine on the horn of T1 and the single basiconic sensillum on A8.
Etymology. The adjectival Latin epithet "sulcata" means "furrowed" and refers to the clearly defined and separate metapleural and paracoxal sulci.
Link Number of basiconic sensilla on A8: one. Shape of male flagellomeres: spherical. Color of head: brown to black. Distal margin of clypeus: serrate. Shape of distal margin of clypeus in anterior view: straight. Width of clypeus: greater than width across toruli. Lateral corner of clypeus: projecting into acute angle. Length of mediofacial striae: not extending above midpoint of compound eye. Anterodorsal node on interantennal process: absent. Central keel: absent. Length of OOL: greater than 2 ocellar diameters. Macrosculpture of frons between median ocellus and inner orbit of eye: absent. Patch of microsculpture posterior to lateral ocellus in male: absent. Patch of microsculpture posterior to lateral ocellus in female: absent. Patch of microsculpture between median and lateral ocelli: absent. Microsculpture on dorsal head: pustulate. Diagnosis. The setose metascutellum of P. taekuli is known to occur in one other species of Paridris, P. spinosa Rajomohana, from India. P. taekuli lacks an occipital carina, has a postmarginal vein about as long as the stigmal vein, and an apically constricted T6. Paridris spinosa has an occipital carina dorsally, a punctiform postmarginal vein and an evenly convex T6.
Etymology. This species is named after Charuwat Taekul, a friend, student and colleague of the authors, for his contributions to the taxonomy and systematics of Platygastroidea.
Link to distribution map. Comments. Specimen OSUC 334142 lacked setae on the metascutellum, as well as on other parts of the body, following the DNA extraction process that we believe caused loss of setation. Because setation of the metascutellum is the most diagnostic character for this species, we exclude the male DNA voucher specimen from the paratype series. The three specimens from the Ivory Coast exhibit notable variability in some of the diagnostic characters, namely, the posterior margin of the metascutellum is slightly emarginate; the setation of the metascutellum is medially reduced; and the postmarginal vein is slightly longer than the stigmal vein wheras it is equal to or less than the length of the stigmal vein in specimens from Asia and Madagascar. Talamas 9,12,20,27,[90][91][92][93][94][95]Morphbank 42 Description. Female body length: 1.56-2.51 mm (n=20). Male body length: 1.26-2.32 mm (n=12).