Masner, a new genus of Ceraphronidae (Hymenoptera, Ceraphronoidea) described using controlled vocabularies

A ceraphronid wasp genus, Masner Mikó & Deans, gen. n., with one species, M. lubomirus Deans & Mikó, sp. n. is described from Australia and Fiji based on male specimens. Th is new genus challenges previous defi nitions of the two extant ceraphronoid families by sharing some character states with Megaspilidae (the presence of an expanded pterostigma and an occipital depression) and other character states with Ceraphronidae (the presence of uniramous anterior protibial spur, presence of a comb of the spur on the proand mesotibial spurs, a single mesotibial spur, an undivided synsternite, the presence of axillular setae, presence of Waterston’s evaporatorium and the fusion of the parossiculus with the gonostipes). Masner is distinguishable from all other Ceraphronoidea by the sensillar patch present on fl agellomere 5 but absent from fl agellomeres 6–9 and the presence of a dorsally visible depression surrounding anterior part of the petiole. Th e limits of Ceraphronidae and Megaspilidae are reviewed, and we provide new characters for family level diagnosis and classifi cation. We also hypothesize that Masner is sister to the remaining


Introduction
Ceraphronoidea is a demonstrably monophyletic lineage (Dowton and Austin 2001, Hymenoptera Tree of Life project in prep., Ronquist et al. 1999), comprised of two moderately sized extant families 1 -Ceraphronidae, with 301 valid species in 14 genera, and Megaspilidae, with 299 valid species in 13 genera (Johnson and Musetti 2009) -and recognized by the fused C+R extending along the anterior margin of the fore wing, the ventrally articulated antennae, the curved fore wing r-rs crossvein, and the presence of two tibial spurs on the fore leg (Dessart and Cancemi 1987;Masner 1993;Westwood 1832).
1 Th e extinct family Stigmaphronidae (and sometimes even Maimetshidae) is also included within Ceraphronoidea (Carpenter 1992, Rasnitsyn 1991) based on the enlarged hind coxa, the relatively ventral articulation of the antennae on the lower face, and the curved fore wing r-rs. We do not treat Stigmaphronidae and Maimetshidae here.
Lubomír Masner played critical roles in ceraphronoid systematics by 1) placing these wasp families into their own superfamily independent of Proctotrupoidea (Masner 1956), and 2) morphologically delimiting the families and subfamilies, in part, by the character states described above (Masner and Dessart 1967), and 3) proposing the main hypothesis of higher-level ceraphronoid relationships (Masner and Dessart 1967, fi g. 40). Th rough his industrious eff orts to collect, sort, mount, label, and re-sort specimens, and by enthusiastically communicating important discoveries, Lubo continues to promote interest in ceraphronoid taxonomy. His encyclopedic knowledge, energy, and willingness to train the next generation of ceraphronoid experts sustain the hope that this fascinating lineage of insects will not wallow in neglect.
Lubo shared one of his discoveries with IM during a recent visit to Ottawa: two lots of specimens that blur the traditional boundaries between Ceraphronidae and Megaspilidae by having only one apical spur on mesotibia while bearing a distinct pterostigma on the fore wing. Lubo has a long history of bringing new taxa to light, instantiated by the descriptions of Pteroceraphron Dessart (Dessart 1981), Retasus Dessart (Dessart 1984), and Aetholagynodes Dessart (Dessart 1994b), as well as the revisions of Cyoceraphron Dessart (Dessart 1994a) and Lagynodes Förster (Dessart 1987b) and descriptions of many new species (e.g., Dessart 1994b) that are based on Masner-sorted CNCI material (see the acknowledgments of these papers). We continue this parade of discovery by describing new taxa and by briefl y discussing the impacts of these observations on our understanding of Ceraphronoidea taxonomy.

Materials and methods
Specimens were borrowed from the South Australian Museum, Adelaide, South Australia (SAMA), and acquired from the collecting eff orts of the Fiji Biodiversity of Arthropods (FBA). Specimens will be deposited in SAMA, the Fiji National Insect Collection, Suva (FNIC), the Canadian National Collection of Insects (CNCI) and the North Carolina State University Insect Museum (NCSU). Digital images were made using an Olympus CX41 compound microscope and DP71 digital camera. SEM micrographs were made using a Hitachi S-3200 Scanning Electron Microscope (wd=23.5, av=5kV). Specimens were critical point dried and coated with palladium prior to examination. For confocal laser scanning microscopy (CLSM) specimens were dissected, and parts of male genitalia were embedded in glycerin gelatin and in type VII agarose. Series of images were taken using a Leica inverted DM IRBE CLSM with a planachromat 40× 1.4 NA oil immersion objective. Videos generated through microscopy are available from the North Carolina State University Insect Museum website and from Google Video; links are in the appendix.
In an eff ort to use unambiguous terms in our taxon descriptions and to make semantic statements about the phenotypes of these taxa we employed several ontolo-gies (i.e., controlled vocabularies) available from the Open Biomedical Ontologies Foundry (OBO Foundry 2 ): Units of Measurement Ontology (UO 3 ; date: 04:03:2009 17:16), Spatial Ontology (BSPO 4 ;date: 08:12:2008 21:42), and the Phenotype and Trait Ontology (PATO 5 ;date: 30:03:2009 16:12). Anatomical terminology follows the Hymenoptera Anatomy Ontology 6 (HAO; Deans et al. 2009) and includes terms from Ronquist and Nordlander 1989, Schulmeister 2001, Mikó et al 2007and Vilhelmsen et al. 2009. New anatomical terms were added to the HAO and are provided below with genus-diff erentia defi nitions (Neuhaus et al. 2004;Smith 2005). We worked to manually compose our descriptions as collections of entity-quality (E-Q) statements, whereby the anatomical structure is the entity and the phenotype descriptor(s) represents the quality -e.g., mesodiscrimen (= entity) foveolate (= quality) (see Mabee et al. 2007a,b for discussion). . anterolateral mesoscutal corner (aem: Fig  1E): Th e corner that is located anterolaterally on the mesoscutum and is adjacent to the dorsal pronotal corner. apical parossiculal seta (asp: Figs 3C, 4A, 6E; =soie subapicale Dessart and Gärdenfors 1985): Th e seta that is located submedially on the apical margin of the parossiculus.  Fig. 1E, 6B): Th e corner that is located submedially on the dorsal margin of the pronotum and that is adjacent to the anterolateral mesoscutal corner. facial pit (fp: Fig. 6A; Bin and Dessart 1983;=fossette frontale Dessart 1966, =fosette scrobale Gärdenfors 1985, =pore facial Dessart 1992): Th e pit that is located on the upper face dorsal to the intertorular carina and corresponds internally to a conical apodeme and not the tentorium. facial sulcus (Dessart 1978): Th e sulcus that arises medially from the ventral margin of the median ocellus and extends towards the intertorular carina [Note: the facial sulcus is usually foveolate in Ceraphronoidea].
frontal scrobe (fdp: Fig. 6D; Mikó et al. 2007; =supraclypeal depression Dessart 1978, =dépression supraclypéale Dessart 2001: Th e scrobe that is located on the upper face.  (Dessart and Gärdenfors 1985): Th e sclerite that is located distally of the gonostipes. interaxillar sulcus (ias: Fig. 1D; =trait axillaire Dessart and Gärdenfors 1985): Th e sulcus that extends medially between the posterior margin of mesoscutum and the anterior margin of mesoscutellum. interocellar pit (Bin and Dessart 1983;=pore interocellaire Dessart 1992): Th e pit that is located on the upper face between lateral ocelli and corresponds internally with a conical apodeme and not the tentorium. interorbital space (=faciale interoculaire Dessart and Gärdenfors 1985): Th e anatomical line that marks the shortest distance between inner orbits. [Note: Th e mesometapleural sulcus represents the border between the mesopleuron and metapleuron. Th e mesometapleural sulcus corresponds to a ridge and could be homologous with the mesepimeral ridge, which is defi nitely part of mesopleuron (Vilhelmsen et al. in press, Mikó et al 2007). In most Hymenoptera the mesopleuron extends past the mesepimeral ridge (not in Cynipoidea), and therefore its posterior margin is not easily delimited by the ridge, making uncertain the exact location of the border between meso-and metapleura based on the ridge and sulcus (=sillon mesopleuro-métopleurale; Dessart 1991, =mesometapleural suture, Vilhelmsen et al. in press).] mesometapleuron: Th e sclerite that is comprised of the fused meso-and metapleura [Note: Th e border between meso-and metapleura is usually indistinct]. longitudinal metacoxal carina (cxc: Fig. 6B): Th e carina that is longitudinal and is located on the metacoxa [Note: In Ceraphronoidea the longitudinal metacoxal carina usually corresponds to rows of elongated setae and delimits a posterior concave area.] metanotal-propodeal sulcus (mps: Fig. 1D, F, 6B; =foveole métanotale, Dessart 1994b, sillon métanotal Dessart 1996: Th e sulcus that extends along the anterior margin of the fused metanotum and propodeum and delimits an anterior narrow, usually diff erently sculptured area. [Note: Th e sulcus could mark the border between metanotum and propodeum or could be homologous with the metanotal trough (Mikó et al. 2007).] Th e area that is concave, is located on the vertex and is limited anteriorly by the preoccipital carina and posteriorly by the occipital carina. preocellar pit (pp: Fig. 6A; Bin and Dessart 1983;=pore preocellaire Dessart 1992): Th e pit that is located on the upper face adjacent the anterior margin of the median ocellus and corresponds internally to a conical apodeme and not the tentorium. scutoscutellar sulcus (sss: Fig. 1D; Mikó et al. 2007;=sillon axillo-scutellaire Dessart 1994b). Th e sulcus that sets off the mesoscutellum from the mesonotum.  Waterston Dessart 1992): Th e area that is located medially on the acro tergite (Packer 2004) of T6 and is modifi ed for releasing glandular products. [Note: Th e Waterston's evaporatorium is a median, concave, calyx-like structure (wo: Fig. 4D, F) surrounded posteriorly by strongly sculptured cuticle in most Ceraphronidae, whereas it is a strongly sculptured area without any concave anterior part in Masner (wo: Fig. 4C, E). Waterston (1923) originally hypothesized, that this cuticle modifi cation has a respiratory function. Later both Ogloblin (1944) and Dessart (1992) assumed that the structure corresponds to an opening of exocrine glands and could have evolved as an increased evaporative surface. Diff erent forms of cuticle modifi cations are known in Hymenoptera around the opening of type III exocrine glands (Noirot and Quennedey 1974, Buckingham and Sharkey 1988, Quicke 1990). We have located glands (g: Fig.  4F) connected to the lateral part of the calyx-like via ducts (d: Fig. 4F) in Aphanogmus and Ceraphron. Ogloblin proposed the term Waterston's organ for the cuticle modification on metasomal tergum 6 described by Waterston (1923): "tergite iv with anteromedianly the remarkable reticulated chitinous ring." Although Ogloblin (1944) suggested that Waterston's structure might have glandular function, he never described any corresponding glands and applied the term "Waterston's organ" exclusively to the cuticle modifi cation. According to the Common Anatomy Reference Ontology (Haendel et al. 2007) a simple organ is a multi-tissue structure. Because this cuticular modifi cation is not a multi-tissue structure we prefer to replace "organ" with "evaporatorium" from the heteropterists' lexicon. Evaporatorium refers to a modifi ed cuticular structure associated with the opening of an exocrine gland, the purpose of which is to enlarge the evaporative surface (Carayon 1962;Torre-Bueno 1989 Diagnosis. Masner gen. n. is distinguishable from all other Ceraphronoidea by three character states: the sensillar patch present on fl agellomere 5 but absent from fl agellomeres 6-9 (sf: Figs 5A, B), dorsally visible depression surrounding anterior part of the petiole and the irregular areolate sculpture of the dorsal cranium. It diff ers from all other Ceraphronidae in presence of pterostigma (sg: Fig 3A) and occipital depression (od: Fig.  1B). Masner gen. n. belongs to Ceraphronidae on the basis of the presence of calcar comb, absence of anterior mesotibial spur, uniramous anterior protibial spur (calcar), absence of anteriorly delimited narrow region of synsternite, presence of axillular setae and parossiculus fused with gonostipes and presence of Waterston's evaporatorium.
Etymology. Th e new genus group name is the Latinized family name of Lubomír Masner, who discovered the unique character combination of Masner gen. n., the presence of pterostigma on the wing and lacking posterior apical mesotibial spur and presorted the specimens of the new genus. Gender is masculine. Color: body brown, (with) low variability of color; maxillo-labial complex yellow; mandible yellow; blade brown; clypeus yellow; radicle yellow; scape yellow; pedicel yellow; leg yellow; tegula yellow; wing base yellow; syntergum yellow; synsternite yellow; male genitalia yellow.
Etymology. Th e dense setae on the clypeus of Masner lubomirus resembles the recumbent pilosity on the chin of Lubomír Masner. Th erefore the new specifi c epithet is the Latinized fi rst name of Lubomír Masner, in the nominative singular case.
Material examined. Holotype male: AUSTRALIA: Queensland, Mount Glorious, 27°19'54"S 152°45'29"E, 7-13. II.1998 Discussion. Ceraphronoidea is divided into two families: Ceraphronidae and Megaspilidae based on ten two-state characters that are, with one exception, invariable within each family (Table 2.). Th e pterostigma is present only in Megaspilinae, but absent in Lagynodinae and Ceraphronidae. Th is observation, and the considera- tion of Bethylidae as the sister group of Ceraphronoidea led Masner and Dessart (1967) to assume that the absence of the pterostigma is the ground plan for the superfamily. Although the subfamily and generic classifi cation of the superfamily is currently considered to be unstable, it is widely accepted that Megaspilidae and Ceraphronidae are two distinct and probably monophyletic taxa. Although the presence of non-overlapping two-state characters strongly support the latter hypothesis, it has never been tested phylogenetically. Overlapping characters in Masner weakens both of the above-mentioned hypotheses (apomorphic pterostigma and the dichotomous Ceraphronoidea). Th e presence of the Waterston's evaporatorium is an important character state that indicates Masner belongs in Ceraphronidae. Th e absence of the anterior calyx-like concave area and the presence only of sculptured cuticle, however, could be considered an ancestral state of Waterston's evaporatorium. Th e structure of Waterston's evaporatorium and the fact that Masner shares two family level characters with Megaspilidae leads us to hypothesize that this taxon is the sister to the remaining Ceraphronidae. Future work will focus on testing this hypothesis using molecular and morphological data.

Acknowledgments
We gratefully acknowledge the assistance of Chuck Mooney (North Carolina State University Analytical Instrumentation Facility) with scanning electron microscopy and Eva Johannes (NCSU-NSCORT Department of Botany, N.C. State University) with confocal laser scanning microscopy. We thank Matt Yoder for his valuable comments on antennal sensillum types, Bob Blinn for comments on the evaporatorium