Ooderella Ashmead,

Gibson, Gary A. P., 2017, Revision of world Ooderella Ashmead (Hymenoptera: Chalcidoidea: Eupelmidae), with description of the first males for the genus, Zootaxa 4289 (1), pp. 1-74: 5-13

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http://doi.org/ 10.5281/zenodo.828791

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Ooderella Ashmead
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Ooderella Ashmead 

Ooderella Ashmead, 1896: 6  (key), 10–11 (description). Type species: Ooderella smithii Ashmead  , by monotypy and original designation.

Diagnosis. FEMALE. Brachypterous, with fore wing usually extending flat over mesosoma to propodeum or base of gaster, though sometimes bent upwards near posterior of mesosoma ( Figs 14, 15View FIGURES 10 – 18); membrane behind venation bare (e.g. Fig. 52View FIGURES 45 – 53) to variably densely setose (e.g. Fig. 7View FIGURES 1 – 9) but with hairlike setae and without hyaline cross-band or anterior and posterior spots with white setae. Head in lateral view variably highly convex (cf. Figs 11View FIGURES 10 – 18, 46View FIGURES 45 – 53, 91View FIGURES 90 – 98); sometimes with ocellocular mark between posterior ocellus and inner orbit (e.g. Figs 47View FIGURES 45 – 53, 157View FIGURES 156 – 164), and sometimes with lighter-colored line or mediolongitudinal sulcus below anterior ocellus on one or both of frons and scrobal depression (e.g. Figs 156View FIGURES 156 – 164, 165View FIGURES 165 – 173). Eye superficially bare but very sparsely microsetose. Antenna with fl1 short compared to fl2, but at least about as long as wide rather than strongly discoidal (e.g. Figs 5View FIGURES 1 – 9, 13View FIGURES 10 – 18). Mandibles with acute tooth ventrally but apical margin above ventral tooth variably broadly truncate to deeply incurved or angularly incised so mandible variably distinctly bidentate (e.g. Fig. 23View FIGURES 19 – 27: insert) to tridentate (e.g. Fig. 5View FIGURES 1 – 9: insert). Pronotal collar transverse to somewhat longer than wide but more or less quadrangular with lateral margins parallel or only slightly anteriorly divergent, with anterolateral corners right-angled (e.g. Figs 3, 6View FIGURES 1 – 9) to more strongly, conelike produced (e.g. Figs 120, 124View FIGURES 118 – 127). Anterior margin of collar abruptly angled relative to inclined neck at least laterally, and dorsally collar divided mediolongitudinally by deep furrow and usually flat even if sometimes inclined on either side of furrow. Mesoscutum with smooth and shiny mesoscutal flange laterally, but otherwise meshlike reticulate at least anterior of posteriorly carinate portion of lateral lobes (except O. platyscapus  where more transversely reticulate-strigose to reticulate-rugulose, Fig. 201View FIGURES 194 – 204); scutellar-axillar complex with scutellum extending to and acutely angled at transscutal articulation. Mesotibia with oblique groove apically between tibial spur and base of tarsus, and with row of dark apical pegs in region over base of tibial spur; mesotarsus with a single row of dark pegs along either side of basal four tarsomeres. Propodeum highly variable in structure. Gaster yellowish to dark brown, though often variably paler to white basally or subbasally; Gt1 distinctly incised medially but subsequent tergites more shallowly emarginate; syntergum narrowed posteriorly to often lighter-colored and/or more translucent and sometimes variably reflexed, posteriorly rounded or angulate bare region (syntergal flange) behind setae (e.g. Figs 9View FIGURES 1 – 9, 18View FIGURES 10 – 18, 204View FIGURES 194 – 204) (except O. ambigua  , see Discussion).

MALE. New World. Head often with ocellocular mark evident as an angulation (e.g. Fig. 82View FIGURES 81 – 89) but only rarely as a lighter-colored line (e.g. Fig. 206View FIGURES 205 – 213) between anterior margin of posterior ocellus and inner orbit; scrobal depression often transversely Π-like above interantennal prominence (e.g. Figs 81View FIGURES 81 – 89, 101View FIGURES 101 – 109), variably sculptured relative to smoother and shinier frons, and usually with mediolongitudinal light-colored sulcus or line of weakness (e.g. Figs 101View FIGURES 101 – 109, 205View FIGURES 205 – 213), but not a circular depression or pit at apex of interantennal prominence (see Gibson 1995, figs 406–408); with uniformly short and sparse setae on gena lateral to malar sulcus. Antenna with torulus about midway between oral margin and anterior ocellus, the ventral margin obviously above lower orbit and distance between ventral margin of torulus and oral margin slightly greater than distance between dorsal margin of torulus and ventral margin of anterior ocellus (e.g. Fig. 101View FIGURES 101 – 109); scape compressed and broadest subbasally to medially, but variably distinctly narrowed apically; pedicel obviously longer than apical width, subtriangular, without row of differentiated setae along ventral length; flagellum conspicuously elongate-slender, filiform with fl1 (anellus) discoidal; funiculars much longer than wide with fl2 about 4× as long as wide and straight (ventral margin not curved) and subsequent funiculars of similar length or only slightly shorter apically, with sparse and comparatively inconspicuous, very short setae much shorter than width of funicle, and either with multiple overlapping rows of longitudinal, convex mps (multiporous plate sensilla) ( Figs 85View FIGURES 81 – 89, 104, 105View FIGURES 101 – 109) (= Reikosiella  group 3 sensu Gibson 1995, fig. 370) or with each mps having long, apically free, seta-like portion so mps superficially appear as comparatively dense setae subappressed to or projecting from surface at acute angle ( Figs 131, 132View FIGURES 128 – 136) (= Reikosiella  group 4 sensu Gibson 1995, fig. 371); clava as slender as preceding flagellomeres and tapered to apex without distinct micropilose sensory region ventrally, at most only very obscurely differentiated into clavomeres by indistinct transverse sutures, and about as long as combined length of apical two funiculars. Mandibles variably distinctly tridentate. Fore wing hyaline or at most slightly infumate ( Fig. 109View FIGURES 101 – 109); not distinctively elongate-slender, at most length only about 2.7× maximum width; dorsally with variably large and distinct speculum behind parastigma beyond basal cell ( Fig. 86View FIGURES 81 – 89), but region often obscured by setae on ventral surface; marginal vein not distinctively long, less than 0.8× width of wing; postmarginal vein obviously longer than stigmal vein, though apex often difficult to distinguish accurately because vein gradually lightened apically. Mesoscutum with deep, distinct notauli; usually quite distinctly meshlike reticulate over at least anterior half (e.g. Figs 87, 88View FIGURES 81 – 89), though sometimes smoother, more coriaceous posteriorly, and small males sometimes almost entirely meshlike coriaceous-reticulate. Metapleuron bare. Propodeum with median carina.

Old World. Males of a single identified species differ from known New World males by the following: torulus obviously closer to oral margin than to anterior ocellus, with ventral margin at level of lower orbit ( Fig. 205View FIGURES 205 – 213); scrobal depression with light-colored line of weakness mediolongitudinally, though more elongate Π-like, extending almost to anterior ocellus ( Fig. 205View FIGURES 205 – 213); flagellum with fl2–fl8 obviously longer than wide and sparsely and inconspicuously setose, but at most only about 2× as long as wide and with elongate mps extending almost entire length of funicular ( Figs 208, 209View FIGURES 205 – 213); clava about equal to combined length of apical three funiculars and with much longer setae than preceding flagellomeres ( Fig. 208View FIGURES 205 – 213: insert).

Hosts. Hosts are known only for O. americana  , which was reared once from eggs of an unidentified species of Orchelimum Serville  ( Orthoptera  : Tettigoniidae  ) (greater meadow katydids) on Daucus carota  L. ( Apiaceae  ) (wild carrot).

Diversity and distribution. Ooderella  is restricted to the Neotropical and Afrotropical regions (Maps 1, 2) except for a single species, O. americana  , in northeastern USA (Map 1A), though it is possible that the northern limits of at least one Neotropical species may extend into southeastern USA (see under O. setosa  ). Within the Neotropical region , species are known from southernmost Mexico ( Fig. 2View FIGURES 1 – 9 A) through Central and South America , but not the islands of the West Indies other than Trinidad (Maps 1B –H, 2A –C). Of the 12 species recorded from the Neotropical region , only 3 are represented by more than 10 specimens  . A total of 7 of the 12 species, 5 uniquely, are known from Costa Rica. This comparatively large number of species from Costa Rica undoubtedly reflects, at least in part, the greater collecting efforts in Costa Rica in modern times and the ready availability of specimens. Within the Afrotropical region, species are currently known only from south of about 10°S (Map 2D), though this likely is an artefact of collecting and specimen availability. Four of the five species recorded from the Afrotropical region are known from less than ten specimens and only one, O. botswanae  , is recorded from more than one country. It therefore seems inevitable that more extensive collecting in both the Neotropical and Afrotropical regions will not only enlarge the known ranges of the currently described species, but result in other, if not many other, new species. 

Generic limits and species relationships. FEMALES. Gibson (1995) provided a comparative description of Ooderella  relative to other eupelmine genera based on females, but this did not include features of what is here described as O. ambigua  . This newly described species is classified in Ooderella  based primarily on the combination of female brachyptery and pronotal structure, though the pronotal collar is unusual in having the anterior margin most distinctly angulate or protuberant anteromedially on either side of the mediolongitudinal furrow ( Figs 12, 15View FIGURES 10 – 18) rather than anterolaterally (e.g. Figs 3View FIGURES 1 – 9, 21View FIGURES 19 – 27, 47View FIGURES 45 – 53). As such, the dorsal surface is shallowly concave ( Fig. 15View FIGURES 10 – 18), whereas in other species it is flat, even if sometimes it is inclined on either side of the furrow. Except for the absence of a transverse line of dark setae on the ridge that delineates the collar from the neck, the pronotal structure of O. ambigua  is quite similar to that of typical Eupelmus  ( Macroneura Dalman  ) females, which was described as character state 15(4) by Gibson (1995). Other Ooderella  females have a pronotal structure described as character state 15(2), and as either substate 2a or 2b depending on whether the anterolateral corners are right-angled (e.g. Figs 3View FIGURES 1 – 9, 21View FIGURES 19 – 27) or more cone-like protuberant (e.g. Figs 47View FIGURES 45 – 53, 120View FIGURES 118 – 127), respectively. These different pronotal structures were hypothesized to have been derived from a strongly transverse pronotum without a distinctly differentiated collar and neck (character 15, state 1), which is possessed by females of many eupelmine genera, including Brasema Cameron  (see figures in Gibson 1995).

Syntergal structure is even more unusual for O. ambigua  within Ooderella  . Except for this species, females of Ooderella  have the syntergum tapered into a posteriorly rounded or angulate region that often is lighter in color and/or translucent, but at least is bare beyond subapical setae whose apices extend only slightly beyond the posterior margin (e.g. Figs 100View FIGURES 99 – 100, 164View FIGURES 156 – 164, 193View FIGURES 185 – 193). The apical region is sometimes also variably abruptly recurved from a more convex or inclined basal region into a narrower and/or horizontal syntergal flange sensu Gibson (1995) (e.g. Figs 164View FIGURES 156 – 164, 184View FIGURES 174 – 184, 204View FIGURES 194 – 204). This type of syntergum (character 39, state 3) was stated as diagnostic for Ooderella  and 16 other eupelmine genera that were hypothesized to form a monophyletic clade based on this shared structure ( Gibson 1995, fig. 410). The syntergum of O. ambigua  differs by being uniformly dark and evenly tapered in one plane to a transverse posterior margin, and with a row of dark setae along the posterior margin that project conspicuously beyond the margin ( Figs 14, 18View FIGURES 10 – 18). This latter structure (character 39, state 1) and setal pattern were hypothesized as the groundplan states of the syntergum for Eupelminae  by Gibson (2017), and the syntergal structures and setal patterns of other Ooderella  as derived states that evolved through secondary development of the cuticle behind the apical-most line of setae characteristic of character state 39(1). The selective pressure for evolving and the functional advantage of having a flanged syntergum are unknown, as is the number of times similar structures may have evolved convergently. However, the putatively plesiomorphic non-flanged structure is possessed by females of some species of several eupelmine genera ( Gibson 1995, table 1), including those of Brasema  (see figures in Gibson 1995), a genus that is by far most highly speciose and diverse in the New World. This latter genus is not defined by any autapomorphies and very possibly is paraphyletic relative to some other recognized genera such as Zaischnopsis Ashmead  and possibly Anastatus Motschulsky  ( Gibson 1995, 2016).

Females of O. ambigua  also have the least reduced wings within the genus. The fore wing ( Figs 14, 15View FIGURES 10 – 18) is right-angle bent near the posterior of the mesosoma, with the basal portion consisting of a setose basal cell behind the costal cell, and the apical portion beyond the bend constituting the setose disc. The wing also retains the typical chalcid vein complex consisting of a submarginal, long marginal, stigmal, and postmarginal vein. The fore wings of other species of Ooderella  extend at most only slightly beyond the base of the gaster and the venation either is reduced to just the submarginal vein or there are at most short remnants of the other veins (see further below). Similarly reduced fore wings to those of O. ambigua  are possessed by females of some other eupelmine genera, including many Eupelmus  ( Macroneura  ) and at least one undescribed Neotropical species of Brasema  . Interestingly, both types of fore wing rudiments occur also in different species of E. ( Macroneura  ), some having the fore wings abruptly bent upwards near the posterior of the mesosoma as in O. ambigua  and some having more severely reduced wings that extend flat over the mesosoma as for females of other species of Ooderella  ( Gibson 1990; Fusu 2017).

Compared to some other species of Ooderella  , females of O. ambigua  also have relatively plesiomorphic head structures, including the head in lateral view being comparatively lowly convex with the anterior surface evenly curved ( Fig. 11View FIGURES 10 – 18), and in frontal view the scrobal depression being bell-shaped because of slightly incurved lateral margins that distinguish distinct parascrobal regions ( Fig. 10View FIGURES 10 – 18) (character 6, state 1). The frontovertex also lacks ocellocular marks (character 9, state 1), even though the frons is comparatively smooth and shiny with only fine coriaceous sculpture ( Figs 10, 12View FIGURES 10 – 18). Finally, the antenna has a tubular, uniformly thick, slightly curved scape (character 5, state 1) and a comparatively short-clavate flagellum ( Fig. 13View FIGURES 10 – 18) as compared to most other Ooderella  . The acropleuron is also completely bare ( Fig. 17View FIGURES 10 – 18) and the mesoscutum lacks distinctly denser regions of white lanceolate setae posteriorly or anteriorly ( Fig. 15View FIGURES 10 – 18). All of these features are hypothesized as likely plesiomorphic compared to other states exhibited by females of some other Ooderella  species. Of uncertain polarity is that in O. ambigua  the mesoscutal lateral lobes are carinately angled along their entire length ( Fig. 15View FIGURES 10 – 18). Except for O.

platyscapus  , other Ooderella  have the mesoscutal lateral lobes carinately angled only within about the posterior third or less. Based on its atypical pronotal and syntergal structures as well as the combination of other putatively plesiomorphic features, O. ambigua  could possibly represent an unusually modified brachypterous species of Brasema  having a secondarily modified pronotal structure that is only superficially similar (convergent) to that characteristic of Ooderella  , and thus be incorrectly classified to genus. However, propodeal structure in O. ambigua  does not support such a hypothesis. Females of Brasema  have a variably long but broad, flat to lowly convex plical region that is differentiated from the callar regions by comparatively obscure subparallel furrows closer to the spiracles than to the midline (character 27, state 2; Gibson 1995, figs 217–221), which was hypothesized as similar to the groundplan structure of the subfamily. Propodeal structure of O. ambigua  is quite different, the propodeum having quite deep, anteriorly convergent depressions along the foramen that differentiate conspicuously convex callar regions from a medially inclined, short and narrow, somewhat bowtie-like plical region ( Fig. 16View FIGURES 10 – 18) (character 27, state 4). Although propodeal structures of other Ooderella  are quite diverse, the structures are variably similar to, and could be derived from, an O. ambigua  -like structure (see further below),

Even excluding O. ambigua  , Ooderella  is unusually diverse in some other features. Both sexes of most eupelmine genera usually have mandibles that can be assigned quite readily to one of two structures—three similar teeth (tridentate: character 1, state 1) or with a single ventroapical tooth and a broad, truncate to shallowly concave dorsoapical margin (bidentate: character 1, state 2a). Different species or even different females of the same species of Ooderella  exhibit variably distinctly bidentate to tridentate mandibles, the difference resulting from how deeply incurved is the apical margin above the acute ventral tooth and whether the emargination is evenly incurved (e.g. Figs 23View FIGURES 19 – 27, 35View FIGURES 35 – 44) or angularly incised to differentiate two more acutely angled teeth (e.g. Fig. 5View FIGURES 1 – 9). Mandibular structure is not clearly visible in most available females of O. ambigua  , but there appears to be only one distinct tooth ventroapically plus a broad, truncate margin dorsoapically that at least sometimes is obscurely differentiated into two blunt teeth by a slight depression or transverse furrow ( Fig. 13View FIGURES 10 – 18: insert). Tridentate mandibles were hypothesized as the groundplan structure for Eupelminae  by Gibson (1995) and bidentate mandibles as apomorphic, and possibly correlated with egg parasitoidism ( Gibson 1995, 2016). Mandibular structure is clearly visible in only one female of O. americana  , a putative egg parasitoid. Each mandible is quite distinctly tridentate because the emargination dorsal to the ventral tooth is angular ( Fig. 5View FIGURES 1 – 9). However, additional females with exposed mandibles are needed to determine whether there is intraspecific variation, similar to some other Ooderella  species.

Ooderella  is also unusual in that females of some species, including O. ambigua  , have the gaster uniformly dark ( Figs 14, 18View FIGURES 10 – 18) (character 42, state 1), whereas in others it is variably paler to white dorsally and/or ventrally, most commonly subbasally (character 42, state 2). This latter, derived state is perhaps most characteristic of females of Anastatus  , although it is also known for at least some species of several other genera ( Gibson 1995, table 1), particularly species with banded fore wings. Smith (1969) provided line drawings and commented on the similarity between females of Tachydromia lilaniensis Smith  ( Diptera  : Empididae  ) and E. (Eupelmus) orthopterae (Risbec)  , both of which have banded fore wings and are found in the same habitat. He suggested mimicry for the similarly banded wings, stating that the “egg-hunting Chalcid and the prey-hunting Empid would tend to exhibit a similar behavioural pattern and the banded wings, folded over the back would protect both in breaking up the insect outline which in their rather vulnerable position on lichen-covered boulders would give protection from would-be predators” ( Smith 1969: 75). Both the abdomen of the empid and the gaster of E. orthopterae  are entirely dark. However, a white region basally to subbasally on the gaster of species with banded wings likely further visually emphasizes ‘breaking up’ of the habitus into two apparent body regions, regardless of how the wings are held at rest. Such a light-colored region appears to be prone to evolve also in brachypterous females. For example, the brachypterous females of E. ( Macroneura  ) much more commonly have the gaster white ventrobasally ( Fusu 2017) than the macropterous females of E. ( Eupelmus  ) ( Gibson & Fusu 2016), as do brachypterous compared to macropterous females of Brasema  (personal observation). This disruptive color pattern in brachypterous females could also provide some protection against predators while exposed on plants or other substrates.

Ooderella  species also have quite diverse female propodeal structures, although as noted above at least all New World species share structures that might be derived from an O. ambigua  -like propodeum. Unstated in the above description of the propodeum of O. ambigua  is that the posterior margin is carinate laterally, but near the midline each carina curves anteriorly, sinuately, to form a short median carina anteriorly. However, there is a triangular region of lighter-colored cuticle between the carina on either side behind the median carina so that the posterior margin is evenly, Π-like incurved ( Fig. 16View FIGURES 10 – 18). Females of the other New World species also have the posterior margin of the propodeum sinuately incurved on either side of the midline to form variably long contiguous margins or a median carina, but without any intervening cuticle posteriorly (e.g. Figs 7View FIGURES 1 – 9, 60View FIGURES 54 – 62, 97View FIGURES 90 – 98). Females of all Afrotropical species have the posterior propodeal margin evenly incurved, and in O. gymnosoma  ( Fig. 183View FIGURES 174 – 184) and O. platyscapus  ( Fig. 202View FIGURES 194 – 204) there is a variably distinct ^-like region of differentiated cuticle posteromedially similar to the posteromedial propodeal region of O. ambigua  . Two of the five Afrotropical species ( O. botswanae  and O. capensis  ) have the foramen incurved almost to the medially emarginate anterior margin such that the plical region is transverse-lunate ( Figs 162View FIGURES 156 – 164, 171View FIGURES 165 – 173), whereas two others ( O. gymnosoma  and O. platyscapus  ) have a somewhat longer but still relatively short plical region ( Figs 183View FIGURES 174 – 184, 202View FIGURES 194 – 204), and O. kenyaensis  has an unusually long propodeum with a median carina ( Fig. 190View FIGURES 185 – 193).

At least one feature could support all New World species except O. ambigua  as a monophyletic lineage within Ooderella  —the presence of dense white setae anteriorly on the acropleuron and usually slightly on the mesopectus anterior of the acropleural sulcus. Females of O. ambigua  ( Fig. 17View FIGURES 10 – 18) and all Afrotropical species have the acropleuron bare posterior to the extensively setose mesopectus, whereas all New World species have setae anteriorly on the acropleuron. The setae are comparatively sparse and inconspicuous in O. flavida  ( Fig. 26View FIGURES 19 – 27), but form a dense tuft posterior and/or dorsal to sparser white setae on the mesopectus in other New World species (e.g. Figs 8View FIGURES 1 – 9, 41View FIGURES 35 – 44, 51View FIGURES 45 – 53). A second feature that could support monophyly of all New World species except O. ambigua  is the presence of a differentiated region of white lanceolate setae on the mesoscutum submedially to posteromedially. The mesoscutum of females of O. ambigua  has mostly inconspicuous, posteriorly directed, hairlike setae except for some anteromedially directed, slender-lanceolate white setae over about the lower half of the inclined inner surfaces of the lateral lobes ( Fig. 15View FIGURES 10 – 18). Females of O. flavida  have the mesoscutum inconspicuously setose with posteriorly directed hairlike setae except for a few posteriorly directed white lanceolate setae posteromedially ( Fig. 24View FIGURES 19 – 27). All other New World species have much more conspicuous, variably elongate-ovate regions of dense, lanceolate white setae posteromedially (e.g. Figs 6View FIGURES 1 – 9, 33View FIGURES 28 – 34, 40View FIGURES 35 – 44). Consequently, both the acropleural and mesoscutal setal patterns could support O. ambigua  as the sister group of O. flavida  + other New World Ooderella  , with the setal patterns of O. ambigua  representing the groundplan states and those of O. flavida  intermediate states in development of denser, more conspicuous tufts of setae on the acropleuron and mesoscutum in other New World species. Further, all New World species have the mesoscutal setal tufts composed of setae that are in a ‘swirled’ pattern, i.e. consisting of anteriorly directed setae anteriorly, at least a few posteriorly directed setae posteriorly, and with a few medially directed setae medially (e.g. Figs 6View FIGURES 1 – 9, 33View FIGURES 28 – 34, 40View FIGURES 35 – 44) except for O. melanosceles  ( Fig. 50View FIGURES 45 – 53), O. spinositegula  ( Fig. 124View FIGURES 118 – 127) and O. thegalea  ( Fig. 152View FIGURES 147 – 155) in which all the setae are directed posteromedially. This latter setal pattern may support the monophyly of these latter three species (see further below).

In addition to having white lanceolate setae posteromedially, several New World species have medially or posteromedially directed white lanceolate setae anteriorly on the mesoscutum on either side of the midline so as to form two, variably closely contiguous setal tufts (e.g. Figs 50View FIGURES 45 – 53, 77View FIGURES 72 – 80, 114View FIGURES 110 – 117). Although females of all Afrotropical species lack as dense regions of lanceolate setae on the mesoscutum as New World species, some do have somewhat denser regions of lanceolate setae anteriorly and posteriorly. The mesoscutum is extensively setose with posteriorly directed hairlike setae in O. gymnosoma  ( Fig. 180View FIGURES 174 – 184) and O. platyscapus  ( Fig. 201View FIGURES 194 – 204). Females of O. kenyaensis  have a similar setal pattern except posteriorly some setae are directed anteriorly ( Figs 188, 189View FIGURES 185 – 193), whereas both O. botswanae  ( Fig. 161View FIGURES 156 – 164) and O. capensis  ( Fig. 170View FIGURES 165 – 173) have noticeably denser regions of white lanceolate setae both posteriorly and anteriorly, with the anterior setae directed posteriorly and uniformly distributed across the sclerite and the posterior setae directed anteriorly. The differences between the setal patterns in New World and Afrotropical species suggest that denser regions of lanceolate setae anteriorly and posteriorly on the mesoscutum evolved independently in the New World and Afrotropical region, though the selective pressure or adaptive function of such setal regions remains unknown. The groundplan state and polarity of the setal patterns in the Old World species are also uncertain, but the shared patterns of O. gymnosoma  and O. platyscapus  , and O. botswanae  and O. capensis  , could support these as species pairs. Further, among New World species, presence of posteromedially to medially directed white setae anteriorly on the mesoscutum could support monophyly of O. melanosceles  + O. setosa  + O. speculifrons  + O. spinositegula  + O. stenoptera  + O. thegalea  , and, possibly, O. americana  and O. smithii  . Females of O. americana  differ from the first six species by having only one or two differentiated white setae that are widely separated on either side of the mesoscutal medial lobe ( Fig. 6View FIGURES 1 – 9). Females of O. smithii  sometimes have several such posteromedially directed setae anteriorly on the mesoscutum, but even when most numerous they do not form as conspicuous regions of differentiated setae as for the first six species, and usually are not evident without correct angle of lighting ( Figs 95, 96View FIGURES 90 – 98). It is possible that the setal patterns of both O. americana  and O. smithii  represent initial stages in the evolution of more distinct setal tufts. However, the setal pattern of O. americana  might more likely be indicated as independently derived from those of the other seven species based on females having a different head structure and sculpture pattern relative to the other species. Females of O. americana  have a comparatively lowly convex, lenticular head ( Figs 3, 4View FIGURES 1 – 9) and a sculptured frons that curves smoothly into the scrobal depression ( Fig. 1View FIGURES 1 – 9) similar to that of O. ambigua  ( Figs 10, 11View FIGURES 10 – 18) (see further below). Females of O. melanosceles  ( Fig. 47View FIGURES 45 – 53), O. microptera  ( Fig. 55View FIGURES 54 – 62), O. setosa  ( Fig. 74View FIGURES 72 – 80), O. smithii  ( Fig. 92View FIGURES 90 – 98), O. speculifrons  ( Fig. 111View FIGURES 110 – 117), O. spinositegula  ( Fig. 120View FIGURES 118 – 127), O. stenoptera  ( Fig. 138View FIGURES 137 – 146), and O. thegalea  ( Fig. 149View FIGURES 147 – 155) have a variably more highly convex head in combination with a smooth and shiny frons that is abruptly differentiated from a strongly sculptured scrobal depression by at least the sculptural difference (e.g. Figs 45View FIGURES 45 – 53, 72View FIGURES 72 – 80, 90View FIGURES 90 – 98). Consequently, head structure/sculpture pattern may support monophyly of these eight species. If so, apparent absence of differentiated setae anteriorly from the mesoscutum of the only known female of O. microptera  and only obscurely differentiated setae in O. smithii  could represent a transformation series indicating the following relationships: O. microptera  + ( O. smithii  + ( O. melanosceles  + O. setosa  + O. speculifrons  + O. spinositegula  + O. stenoptera  + O. thegalea  )).

As noted above, O. ambigua  has the putative groundplan head states for Ooderella  . Females of O. americana  ( Fig. 1View FIGURES 1 – 9), O. flavida  ( Fig. 19View FIGURES 19 – 27), O. hansoni  ( Fig. 28View FIGURES 28 – 34), O. hyalipleura  ( Fig. 35View FIGURES 35 – 44) and O. reticulifrons  ( Fig. 63View FIGURES 63 – 71) all have the frons, at least in part, similarly sculptured as the scrobal depression, unlike the other eight New World species listed above with a smooth and shiny frons. The frons is entirely, similarly sculptured as the scrobal depression in all five of the above listed species except for O. hyalipleura  , in which it is only partly sculptured in a longitudinal band below the median ocellus ( Fig. 35View FIGURES 35 – 44). Females of O. flavida  also differ from the other species by having a more highly convex head in combination with a scrobal depression that is deep ventrally and shallowed dorsally toward the anterior ocellus. Further, the scrobal depression is abruptly divided by a transverse depression somewhat below mid-height, which differentiates below it a distinct interantennal prominence and separate scrobe above each torulus ( Fig. 19View FIGURES 19 – 27). This scrobal depression structure might represent an independent transformation from an O. ambigua  -like structure or one that is intermediate in development of similar structures shared by O. americana  ( Fig. 1View FIGURES 1 – 9), O. hansoni  ( Fig. 28View FIGURES 28 – 34), O. hyalipleura  ( Fig. 35View FIGURES 35 – 44) and O. reticulifrons  ( Fig. 63View FIGURES 63 – 71). Females of the latter four species all have a strongly, transversely Π-shaped scrobal depression that has a variably distinct transverse line of demarcation separating the scrobes and interantennal prominence below it from a reticulate and strongly though poorly differentiated dorsal portion of the scrobal depression above it. The demarcation line is most noticeable in O. reticulifrons  where it is evident as quite a distinct transverse furrow above each scrobe ( Fig. 63View FIGURES 63 – 71) similar to O. flavida  ( Fig. 19View FIGURES 19 – 27), though this is less developed in the other three species ( Figs 1View FIGURES 1 – 9, 28View FIGURES 28 – 34, 35View FIGURES 35 – 44).

Excluding O. ambigua  , females of O. americana  , O. flavida  , O. hansoni  , O. hyalipleura  and O. reticulifrons  also have fore wings with the venation reduced to just the submarginal vein or at most the submarginal vein and a short remnant of the marginal/postmarginal veins apically (see Remarks section for O. flavida  ), plus a costal cell and at most a very short membranous region beyond the costal cell. Available females of O. flavida  appear to have the membrane behind the venation bare except for marginal setae in the apical half, whereas the membrane is sparsely setose in O. hansoni  ( Fig. 32View FIGURES 28 – 34), O. hyalipleura  ( Fig. 42View FIGURES 35 – 44) and O. reticulifrons  ( Fig. 69View FIGURES 63 – 71) and densely setose in O. americana  ( Fig. 7View FIGURES 1 – 9), but without any apparent discal region. Except for O. ambigua  ( Figs 14, 18View FIGURES 10 – 18), females of the above species also have the gaster variably paler basally or subbasally dorsally and/or ventrally. Females of O. americana  have the gaster paler basally but still brownish to yellowish-brown ( Figs 4, 9View FIGURES 1 – 9), whereas O. flavida  uniquely has the gaster more or less unicolorous dorsally but almost entirely white ventrally ( Fig. 22View FIGURES 19 – 27: insert). Females of O. hansoni  ( Fig. 34View FIGURES 28 – 34), O. hyalipleura  ( Figs 41, 43View FIGURES 35 – 44) and O. reticulifrons  ( Figs 67, 71View FIGURES 63 – 71) have the gaster dorsally whitish subbasally as well as ventrobasally, which might support the monophyly of these three species, though O. speculifrons  also has a gaster that is white ventrobasally ( Figs 112, 117View FIGURES 110 – 117). Females of O. hansoni  ( Fig. 30View FIGURES 28 – 34), O. hyalipleura  ( Fig. 41View FIGURES 35 – 44) and O. reticulifrons  ( Fig. 67View FIGURES 63 – 71) also share long ovipositor sheaths, though the sheaths project quite conspicuously also in O. flavida  ( Fig. 22View FIGURES 19 – 27). Females of the former three species also have a propodeum that is only comparatively sparsely setose with slender, more hairlike setae posterolaterally. Although a relative feature, other species, including O. ambigua  ( Fig. 16View FIGURES 10 – 18), have more conspicuous, slender-lanceolate white setae that often form quite a dense region posterior to the spiracle.

Relationships among the group of eight New World species united by a smooth and shiny frons plus a comparatively highly convex head ( O. melanosceles  , O. microptera  , O. setosa  , O. smithii  , O. speculifrons  , O.

spinositegula  , O. stenoptera  and O. thegalea  ) remain largely unresolved. All share one putative symplesiomorphy—ovipositor sheaths that project only slightly beyond the syntergum, and except for O. speculifrons  one other putative symplesiomorphy—an entirely dark gaster. All except for O. setosa  ( Fig. 78View FIGURES 72 – 80) also share one putative apomorphy—a fore wing having the membrane behind the submarginal vein bare. As discussed above, differences in development of differentiated setae anteriorly on the mesoscutum could indicate O. microptera  and O. smithii  represent basal clades of the group. However, females of O. smithii  , O. melanosceles  , O. spinositegula  and O. thegalea  have slightly different head shapes than those of O. microptera  , O. speculifrons  and O. stenoptera  . In lateral view, the first four species have a somewhat more highly convex head such that this is broadest dorsally ( Figs 46View FIGURES 45 – 53, 91View FIGURES 90 – 98, 119View FIGURES 118 – 127, 148View FIGURES 147 – 155), whereas the last three species have the head broadest within the ventral half or close to the middle ( Figs 55View FIGURES 54 – 62, 112View FIGURES 110 – 117, 140View FIGURES 137 – 146). Regardless, O. melanosceles  , O. spinositegula  and O. thegalea  almost certainly represent a monophyletic lineage. As noted above, this is supported by females of the three species having the posterior mesoscutal setal tuft composed entirely of posteromedially directed setae. It is also supported by females having more strongly, cone-like produced pronotal corners ( Figs 47View FIGURES 45 – 53, 124View FIGURES 118 – 127, 152View FIGURES 147 – 155) than those of the other species, an only coriaceous to coriaceous-imbricate rather than reticulate scrobal depression, and distinct tegular tufts of setae ( Figs 51View FIGURES 45 – 53, 126View FIGURES 118 – 127, 155View FIGURES 147 – 155), although females of O. americana  also have quite distinct tegular setal tufts ( Figs 6, 8View FIGURES 1 – 9) and the setae are obviously denser along the inner margin posteriorly in some other species. Monophyly of the three species is likely further supported by their longer fore wings and ventrally bare costal cell. Unlike other New World Ooderella  females with fore wings extending only to about the base of the gaster or less (i.e. excluding O. ambigua  ), the submarginal vein extends at most about two-thirds the length of the wing and there is a comparatively long, narrower membranous region extending beyond the costal cell ( Figs 52View FIGURES 45 – 53, 125View FIGURES 118 – 127, 153View FIGURES 147 – 155). The absence of any vein remnant in this membranous portion beyond the submarginal vein, unlike for O. gymnosoma  (see below), could indicate the apical membranous region is secondarily derived. If so, presence of the longest fore wings as well as quite distinct ocellocular marks in O. melanosceles  ( Fig. 47View FIGURES 45 – 53) and at least larger O. spinositegula  ( Fig. 120View FIGURES 118 – 127) females support a O. thegalea  + ( O. melanosceles  + O. spinositegula  ) relationship, although females of O. smithii  also have quite distinct ocellocular marks ( Fig. 92View FIGURES 90 – 98). Further, Afrotropical females have ocellocular marks whether or not the frons is smooth and shiny or distinctly sculptured ( Figs 157View FIGURES 156 – 164, 167View FIGURES 165 – 173, 176View FIGURES 174 – 184, 186View FIGURES 185 – 193, 196View FIGURES 194 – 204).

Monophyly of the five recognized Afrotropical species may be supported by females sharing a similar syntergal structure in which one or a few transverse rows of setae differentiate a bare, convexly inclined basal region from a more horizontal, bare syntergal flange ( Figs 164View FIGURES 156 – 164, 184View FIGURES 174 – 184, 193View FIGURES 185 – 193, 204View FIGURES 194 – 204). All Afrotropical females also share a conspicuously highly convex head ( Figs 158View FIGURES 156 – 164, 166View FIGURES 165 – 173, 175View FIGURES 174 – 184, 187View FIGURES 185 – 193, 195View FIGURES 194 – 204) whether or not the frons is distinctly sculptured. This head structure must be convergent to that of some New World species if the New World species are monophyletic. Further, the gaster is variably paler subbasally, dorsally and/or laterally ( Figs 164View FIGURES 156 – 164, 168View FIGURES 165 – 173, 182View FIGURES 174 – 184, 193View FIGURES 185 – 193, 204View FIGURES 194 – 204) in all Afrotropical females except for some specimens of O. capensis  . Again, similar subbasally pale gastral color patterns in some New World species must be convergent if the New World and Afrotropical species each represent monophyletic lineages. The ovipositor sheaths are short and the fore wings are setose in all Afrotropical species except O. gymnosoma  ( Figs 177, 183View FIGURES 174 – 184), though the former two states likely are symplesiomorphic and the bare fore wing membrane and long ovipositor sheaths of O. gymnosoma  likely convergent to similar states in some New World species. As noted above, the different mesoscutal setal patterns of females could support O. botswanae  + O. capensis  and O. gymnosoma  + O. platyscapus  as species pairs, though if the setal pattern of the last two species represent the groundplan state the different setal patterns would indicate the following relationship: O. gymnosoma  and/or O. platyscapus  + ( O. kenyaensis  + ( O. botswanae  + O. capensis  )). The similarly reduced wing venation clearly not extending to the apical margin of the wing shared by O. botswanae  and O. capensis  females could further support their sister-group relationship. Females of the three other Afrotropical species have the venation extending to the apical wing margin with a variably distinctly differentiated apical portion possibly representing a remnant of the marginal/postmarginal veins or, in the case of O. gymnosoma  , with the submarginal vein clearly not extending to the wing margin but with a detached remnant of the marginal/postmarginal veins still evident apically. Females of O. botswanae  and O. capensis  , and of O. gymnosoma  and O. platyscapus  , also share similar propodeal structures that could support the four species as two species pairs, but the groundplan propodeal structure and therefore likely structural transformation series remain uncertain.

Females of O. botswanae  have a sulcus or light-colored line extending mediolongitudinally on the frons below the anterior ocellus into the scrobal depression ( Figs 156, 157View FIGURES 156 – 164) in addition to distinct ocellocular marks ( Fig. 157View FIGURES 156 – 164). Females of the Afrotropical genus Cervicosus Gibson  have a similar head structure, including being highly convex with an even more distinct mediolongitudinal sulcus ( Gibson 1995, character 10, state 2; fig. 451) and ocellocular marks, though females are perhaps at least superficially more similar to females of O. platyscapus  in having the scape conspicuously compressed, a more finely sculptured mesoscutum, and a comparatively long pronotum. In addition to having an elongate-pentagonal pronotum with subparallel sides, Cervicosus  females are also brachypterous and have a strongly transverse propodeum that most closely resembles the propodea of O. ambigua  and O. capensis  . Unlike for Ooderella  species, the dorsal surface of the pronotum is essentially in one plane without a distinctly differentiated collar and neck. Gibson (1995) distinguished two species in Cervicosus  and although he hypothesized the two constituted a monophyletic lineage he also postulated that recognition of Cervicosus  might render Ooderella  paraphyletic ( Gibson 1995, fig. 519), simply representing two Afrotropical species in which the pronotum was secondarily ‘flattened’ in the common ancestor so that the abruptly differentiated collar and neck diagnostic of Ooderella  species was lost. Another difference between species of the two genera is that fl1 is more strongly reduced, anelliform, in Cervicosus  . The two genera could each be monophyletic sister-genera if the longer fl1 characteristic of Ooderella  is secondarily derived, but unfortunately the present study provides no new information in resolving relationships between these two genera or other Eupelminae  .

MALES. Even though males were unknown for the genus, Gibson (1995, fig. 519) included Ooderella  within a clade of genera supported by four hypothesized synapomorphies, two of which based on males—ventral margin of torulus approximately in line with lower orbit (character 48, state 2), and flagellum filiform with both setae and mps short and dense, and the mps in multiple rows per segment (character 50, state 2b). However, New World males here interpreted as Ooderella  have the toruli quite obviously above the lower orbit (character 48, state 1) (e.g. Figs 101View FIGURES 101 – 109, 128View FIGURES 128 – 136), though this is affected by differences in the angle at which the head is viewed ( Fig. 81View FIGURES 81 – 89). Both New and Old World males also have a flagellum with very short and sparse setae, and either dense mps that are quite short and in multiple rows along each funicular ( Figs 85View FIGURES 81 – 89, 105View FIGURES 101 – 109, 209View FIGURES 205 – 213) (character 50, state 2c; fig. 370) or with a long, apically free portion such that each mps superficially appears as an apically directed seta ( Fig. 132View FIGURES 128 – 136) (character 50, state 2d; fig. 371). These two latter flagellar types are characteristic of what Gibson (1995) keyed, respectively, as males of “? Reikosiella  group 3” (couplet 40) and “? Reikosiella  group 4” (couplet 39). Gibson (2017) later synonymized Reikosiella  under Merostenus Walker  and considered true Merostenus  males to have a flagellum with variably sparse but comparatively long setae at least about as long as the width of the funicular, and if with dense mps then these long and not in multiple rows per funicular. The above differentiation of Ooderella  males from those of Merostenus  is based on some males with type 3 and type 4 flagella being associated with females through identical label data, as well as the males sharing some morphological features with females. Although mesoscutal sculpture is variable, most New World males here identified as Ooderella  have quite a coarsely meshlike reticulate mesoscutum, which is also characteristic of Ooderella  females. Fusu (2013) described the males of three Palearctic species of Merostenus  as imbricate-coriaceous and Gibson (2017) described males of some other Old Word Merostenus  species as being variably more strongly reticulate in part. Some small individuals I identify as Ooderella  males have quite a finely sculptured mesoscutum, but at least New World males I identify as Merostenus  have the mesoscutum meshlike coriaceous to coriaceous-imbricate. Also, some of the males I identify as Ooderella  have a transversely Π-like scrobal depression that resembles the scrobal depression of some females with identical collections records, and most have a vertical line of weakness or light-colored sulcus within the scrobal depression (character 47, state 2; Figs 81View FIGURES 81 – 89, 101View FIGURES 101 – 109, 128View FIGURES 128 – 136, 205View FIGURES 205 – 213). Males identified as Merostenus  often have a small depression or pit at the apex of the interantennal prominence ( Gibson 1995, figs 407, 408; Fusu 2013, fig. 43), but not a vertical sulcus or lighter-colored line within the scrobal depression. However, doubt remains concerning generic identity of males identified as Ooderella  versus Merostenus  because some New World males identified as one or the other genus have similar fore wing setal patterns. Some males assigned to both genera have a dorsally bare region (speculum) on the fore wing disc delineated basally by the basal setal line and posteriorly by the mediocubital setal line, though often the bare region is obscured by setae on the ventral surface. More reliable differentiation of males of Ooderella  and Merostenus  requires definitive association of the sexes through rearing or, more practicable and feasible, through COI analysis.

Those males of Ooderella  having a flagellum with short mps in multiple overlapping rows and very short setae (e.g. Fig. 105View FIGURES 101 – 109) can be mistaken for males of Anastatus  ( Gibson 1995, fig. 362) or those Zaischnopsis  males with a similar flagellar structure that Gibson (1995, fig. 361) differentiated within the genus as “flagellar type 1”. Anastatus  males always have bidentate mandibles whereas Zaischnopsis  males have distinctly tridentate mandibles, though because of variation in Ooderella  this is not necessarily helpful in correctly identifying Ooderella  males. However, typical Anastatus  and Zaischnopsis  males differ from Ooderella  males in having fl2 curved in lateral view or at least having the ventral surface slightly concave ( Gibson 1995, figs 361, 362). Further, such New World males have the toruli inserted obviously closer to the oral margin than the anterior ocellus at or somewhat below the lower orbits.

I have seen 43 males from the Neotropical region ( Bolivia, Brazil, Costa Rica, Ecuador, Peru, Trinidad and Venezuela) that I distinguish as Ooderella  males. Of these, I assign 21 to three species— O. setosa  , O. smithii  and O. spinositegula  , based primarily on the males being co-collected with females. Of the remaining 21 (all CNC) not assigned to species, 3 have flagellar structures similar to that described for O. spinositegula  , whereas the other 19 have flagellar structures similar to that described for O. setosa  and O. smithii  . Several are most similar to males described as O. setosa  and might be this species, but they are not included in the type series because they were not collected with females and mostly are from different localities than females. Although I have not seen any female Ooderella  from southern USA, two males ( CNC) from Florida are very similar to O. setosa  males. Because the latter species is known from Trinidad as well as Central America it is quite possible that O. setosa  occurs also in subtropical Florida (see under this species). Males are known for only one of the species described from the Afrotropical region . They are similar to New World males assigned to Ooderella  in having very short and sparse flagellar setae, but differ in having much shorter funiculars ( Fig. 208View FIGURES 205 – 213) and longer mps extending most of the length of the funicular ( Fig. 209View FIGURES 205 – 213). However, the two known males do have a yellowish line between the interantennal prominence and anterior ocellus ( Fig. 205View FIGURES 205 – 213), which supports an association with New World Ooderella  . Relatively distinct ocellocular marks ( Fig. 206View FIGURES 205 – 213) also supports association of the sexes. 

COI

University of Coimbra Botany Department

CNC

Canadian National Collection of Insects, Arachnids, and Nematodes

Kingdom

Animalia

Phylum

Arthropoda

Class

Insecta

Order

Hymenoptera

Family

Eupelmidae

Loc

Ooderella Ashmead

Gibson, Gary A. P. 2017
2017
Loc

Ooderella

Ashmead 1896: 6