Dipara Walker, 1833

Desjardins, Christopher A., 2007, Phylogenetics and classification of the world genera of Diparinae (Hymenoptera: Pteromalidae), Zootaxa 1647 (1), pp. 1-88 : 42-51

Dipara Walker

Dipara Walker 1833: 371 , 373. Type species: Dipara petiolata Walker (by monotypy).

Tricoryphus Förster 1856 . Type Species: Tricoryphus fasciatus Thomson (by subsequent monotypy ( Thomson 1876)) [Synonymized by Domenichini 1953]

Apterolelaps Ashmead 1901 . Type Species: Apterolelaps nigriceps Ashmead (orig. desig. and by monotypy) [Synonymized by Graham 1969]

Alloterra Kieffer and Marshall 1904: 46–47 . Type species: Alloterra claviger Kieffer and Marshall (by monotypy). New synonymy. [Type specimen of genus not examined]

Trimicrops Kieffer 1906 . Type Species: Trimicrops claviger Kieffer (by monotypy). New synonymy. [Type specimen of genus not examined]

Parurios Girault 1913 [175]: 318. Type species: Parurios australiana Girault (by monotypy). New synonymy. [Type specimen of genus not examined]

Epilelaps Girault 1915 [232]: 344. Type species Epilelaps hyalinipennis Girault (orig. desig.). [Synonymized by Bouček 1988]

Pseudipara Girault 1915 [232]: 345. Type species: Pseudipara albiclava Girault (orig. desig. and by monotypy). New synonymy. [Type specimen of genus examined]

Uriolelaps Girault 1915 [239]: 201. Type species: Uriolelaps argenticoxae Girault (orig. desig.) New synonymy. [Type specimen of genus not examined]

Hispanolelaps Mercet 1927 . [Synonymized by Domenichini 1953]

Pseudiparella Girault 1927 [416]: 334–335. Type species Pseudiparella emersoni Girault (by monotypy). [Synonymized by Bouček 1988]

Emersonia Girault 1933 [441]: [1]. Type species: Emersonia atriscutum Girault (by monotypy). New synonymy. [Type specimen of genus not examined]

Grahamisia Delucchi 1962: 379–380 . Type species: Grahamisia saetosa Delucchi (orig. desig. and by monotypy). New synonymy. [Type specimen of genus not examined]

Afrolelaps Hedqvist 1963: 47 . Type species: Afrolelaps maculata Hedqvist (orig. desig.). New synonymy. [Type specimen of genus not examined]

Pondia Hedqvist 1969: 197 . Type species: Pondia punctulata Hedqvist (orig. desig.). New synonymy. [Type specimen of genus examined]

Diparomorpha Hedqvist 1971: 57–58 . Type species: Diparomorpha machadoi Hedqvist (orig. desig. and by monotypy). New synonymy. [Type specimen of genus not examined]

Diagnosis: Dipara females can be identified by a combination of features. First, at least one pair of setae (or

bristles; Fig. 53 ) are present on the lateral margins of the petiole. The only other females with these setae are Chimaerolelaps , Lelaps , and Neapterolelaps . Dipara is distinguished from Chimaerolelaps by an anellus that is broader than long and at most 2 pairs of scutellar bristles. Chimaerolelaps has an anellus that is longer than broad and 4 pairs of scutellar bristles. Dipara can be distinguished from Lelaps by the absence of a median clypeal tooth, which all species of Lelaps have. Finally, Dipara can be easily distinguished from Neapterolelaps by the characteristics discussed in the diagnosis for Neapterolelaps . Dipara males can be separated from Lelaps males by their lack of a median clypeal tooth. They can be distinguished from Netomocera males by their elongate petiole (>2X longer than broad), whereas Netomocera males have a petiole that is broader than long.

Discussion: Tricoryphus Förster and Hispanolelaps Mercet were synonymized with Dipara by Domenichini (1953). Apterolelaps Ashmead was synonymized by Delucchi (1958) with Tricoryphus Förster and then with Dipara Walker by Graham (1969). However, both Hedqvist (1969) and Yoshimoto (1977) treated Apterolelaps as a valid genus based on the absence of an anellus. Heydon and Bouček (1992) resynonymized Apterolelaps , suggesting that the type specimen was aberrant in having a partially fused anellus. Heydon and Bouček then stated that Yoshimoto’s (1977) description of Dipara pedunculata matched Apterolelaps , although the type specimen he selected differed from the description and appeared identical to Dipara canadensis Hedqvist. Heydon and Bouček therefore synonymized Dipara pedunculata Yoshimoto with Dipara canadensis Hedqvist.

Dipara sensu Bouček (1988) and groups of taxa which appeared intermediate to Dipara and Parurios were divided into multiple taxonomic units for the phylogenetic analysis, which are discussed in the Taxonomic Scope section. In the following discussion, these taxonomic units are referred to in angle brackets, so as not to mistake them for valid names. The following genera rendered Dipara sensu Bouček (1988) paraphyletic in all phylogenetic analyses and are herein synonymized with Dipara : Afrolelaps Hedqvist , Alloterra Keiffer and Marshall , Emersonia Girault , Grahamisia Delucchi , Parurios Girault , Pondia Hedqvist , Pseudipara Girault , Trimicrops Keiffer , and Uriolelaps Girault. Within this revision, the reasoning behind synonymy is generally given before the synonymy itself. However, the situation with Dipara is so complex that it is important to provide an overview before examining the specific details.

Grahamisia Delucchi is extremely similar to Dipara s. s., and in many museum collections the former is listed as a synonym of the latter, although the two have never been formally synonymized. Bouček (1988) maintained Grahamisia as a valid genus, although he stated that further examination of the African fauna may lead to its synonymy. Grahamisia and Dipara s. s. group together in all analyses based on 2 synapomorphies: a laterally bulging pronotum and a heavily sculptured mesepimeron. Grahamisia differs from Dipara s. s. only in notaular structure and the presence of black circular markings on the lateral lobes of the scutum. Grahamisia also renders Dipara sensu Bouček (1988) paraphyletic in the phylogenetic analysis, and for these reasons Grahamisia and its synonym Afrolelaps are synonymized with Dipara .

Bouček (1988) maintained Pseudipara Girault as a genus, presumably for two reasons. First, it has a more slender habitus than Dipara , and second, it has a much longer petiole. In the phylogenetic analysis Pseudipara renders Dipara sensu Bouček (1988) paraphyletic, and generally groups with Alloterra and <Micro Dipara > based on having notauli which meet the posterior scutal margin close together but are not strongly arched. Due to the lack of qualifiable morphological differences between Pseudipara and Dipara and the phylogenetic positioning of Pseudipara , Pseudipara is synonymized with Dipara .

Additionally, both Parurios Girault and Alloterra Kieffer and Marshall rendered Dipara sensu Bouček (1988) paraphyletic in the phylogenetic analysis. Unlike the majority of diparine genera, Dipara sensu Bouček (1988) and Parurios have traditionally been separated by male morphology. In the majority of keys (e.g., Bouček 1988), the males of both Alloterra and Parurios are identified by their short, cylindrical flagellar segments with short setae while Dipara s. s. has long, pedunculate segments with long setae. Since Alloterra and Parurios have not been thought to overlap in distribution, and since the antennal structure has been thought to be a reliable separator of Dipara sensu Bouček (1988) and Parurios , this has seemed effective. However, both Alloterra and Parurios are present in Central America, and this had lead to the misidentification of many neotropical Parurios males as Alloterra and Parurios females as Dipara . Thus, the male of Alloterra is indistinguishable from the male of Parurios , and other than geographically the two genera have limited overlap. Also, a diparine species from Tanzania was examined that appears to have typical Dipara s. s. -like females and typical Parurios -like males. Although the specimens were not reared, and therefore no certain association can be made, the simultaneous collection of a large series of both sexes and strong similarities in mesosomal sculpture suggest they are conspecific. Therefore, the cylindrical antennae may not be as reliable an indicator as previously thought.

Alloterra and Parurios are not resolved as sister-taxa in any of the phylogenetic analyses. Additionally, molecular evidence (Desjardins et al. unpublished) supports a sister-group relationship between Alloterra and <Australian Dipara > rather than one between Alloterra and Paurios. In the molecular analysis, Alloterra and <Australian Dipara > are shown as sister-groups, with Parurios sister to that clade, and have much higher sequence similarity to each other than either does to Parurios . While this information in itself could suggest that the pedunculate antennae of Dipara is derived relative to the cylindrical antennae of Alloterra and Parurios , character state reconstruction of the morphological data shows that Dipara -like antennae are the ancestral state for the entire clade.

Bouček (1988) separated the females of Dipara from Parurios based on the position of the median scutal bristles. However, he noted that this character system only worked with the Australian fauna, as non-Australian Dipara would be identified as Parurios in his key. No character was found that would reliably separate female Parurios from Dipara sensu Bouček (1988) at the world level, and if antennal morphology is as plastic as the previous discussion suggests, there is no reliable way to separate these groups in either sex. For these reasons, Parurios and its synonyms, Emersonia and Uriolelaps , are synonymized with Dipara .

The female of Alloterra Kieffer and Marshall is a highly modified, minute, apterous diparine with reduced eyes and no ocelli. However, as previously mentioned, the phylogenetic analysis supports its position within Dipara sensu Bouček (1988) , and the male of Alloterra is indistinguishable from Parurios -like males. (It should be noted that Yoshimoto (1977) keyed the male of Alloterra based on the presence of a spur on the posterior surface of the metacoxa. However, this spur is actually an extension of the lower-most metacoxal striation, and occurs to varying degrees throughout both Dipara sensu Bouček (1988) and Parurios -like males.) For the aforementioned reasons, Alloterra and its synonym Trimicrops are herein synonymized with Dipara .

Pondia Hedqvist either is nested within a monophyletic Dipara (3 phylogenetic analyses) or lies at the base of the clade containing the most derived diparines (1 phylogenetic analysis). When in the latter position, the clade of Pondia + the derived diparines is supported by 3 synapomorphies: a reduced prepectus, reduced axillae, and the loss of the median scutal bristles. The reduction of the prepectus and axillae may play a functional role in the further specialization of diparines to litter habitats; none of the diparines within clade inclusive of Pondia have winged females (or even brachypterous females). Regardless, Pondia Hedqvist renders Dipara paraphyletic in all analyses and the former herein synonymized with the latter.

Diparomorpha Hedqvist is the one described diparine genus not included in the phylogenetic analysis, as it is known only from the type specimen which could not be located. Hedqvist denoted a variety of depositories for his types throughout his descriptions, including his own personal collection. However, most of his holotypes are housed at the British Museum of Natural History, regardless of the listed type depository. Two holotypes were acquired through Christer Hanssen (Swedish Museum), Diparisca ferriei and Dipara canadensis , which were stated to be the only types remaining in his personal collection. The holotype of Diparomorpha (which is known only from the type) is located neither in the BMNH nor in Hedqvist’s personal collection. Furthermore, contact could not be established with the Laboratório de Biologia, Dundo, Lunda, Angola, which is the listed depository for the specimen.

From the description and illustrations, it is apparent that Diparomorpha belongs in Dipara . The former differs from the latter in only two respects. Although Heqvist describes Diparomorpha as having 7 funicular segments and no anellus, his illustration suggests that an anellus is present. Diparomorpha also completely lacks notauli (which appeared to be Hedqvist’s justification for describing it as a new genus), which it shares only with Pyramidophoriella . However, notaular form is extremely variable in Dipara and more broadly in Diparinae , and this likely represents an autapomorphic state within Dipara . In addition, Diparomorpha possesses an extremely dorsally flattened thorax, and although an illustration in dorsal view is not provided, the description is similar to the thorax of Dipara turneri and the taxonomic unit <Fijian Dipara / Parurios >. As both the latter taxa have very fine and weakly impressed notauli, it is possible that notauli were lost entirely by Diparomorpha within this clade. Therefore Diparomorpha is also synonymized with Dipara , and likely occupies a phylogenetic position near the aforementioned taxa.

Number of Species: 39 described species, possibly hundreds of undescribed species.

Distribution: Cosmopolitan.

Hosts: An undescribed Indian species ( Parurios Girault ) was reared from a curculionid ( Coleoptera ) feeding on the roots of Cyperus ( Bouček 1988) .

Key to species: None.

Figures

FIGURES 53–58. Pondia sp.: 53, mesosoma (dorsal view). Pyramidophoriella sp.: 54, mesosoma (lateral view); 55, mesosoma (dorso-lateral view). Cerodipara sabensis: 56, head (frontal view); 57, head (latero-frontal view); 58, mesosoma (dorsal view).