Penthetria Meigen

• Main Author: Fitzgerald, Scott J.
• Format: Text
• Language: unknown
• Published: Zenodo 2021
• Subjects: Biodiversity; Taxonomy; Animalia; Arthropoda; Insecta; Diptera; Bibionidae; Penthetria; Canada; Pacific; Indian; Bergen; British Columbia; Stump; Fitzgerald; Heer; Brune; Sakhalin; ren
• Online Access: https://dx.doi.org/10.5281/zenodo.4558063; https://zenodo.org/record/4558063

Genus Penthetria Meigen Amasia Meigen, 1800: 20. Unavailable name; work suppressed for purposes of zoological nomenclature by I.C.Z.N., 1963: 339. Penthetria Meigen, 1803: 264. Type species: Penthetria funebris Meigen, 1804: 104 (by monotypy in Meigen 1804: 104). Threneste Wiedemann, 1830: 618. Nomen nudum . Eupeitenus Macquart, 1838: 88 (also 1838: 84). Type species: Penthetria atra Macquart, 1834: 175 (by monotypy), [examined; BMNH]. Crapitula Gimmerthal, 1845: 330. Type species: Crapitula motschulskii Gimmerthal 1845: 330 (by monotypy). * Bibiopsis Heer, 1849: 228. Type species: Bibiopsis cimicoides Heer, 1849: 229 (designated by Carpenter 1992: 414). * Mycetophaetus Scudder, 1892: 20. Type species: Mycetophaetus intermedius Scudder, 1892: 20 (by monotypy), [examined; MCZC]. [synonymized by Fitzgerald 1999]. Amasia Meigen in Hendel, 1908: 50. Type species: Penthetria funebris Meigen as “ Amasia funebris (Meig.),” by subsequent monotypy in Bezzi, 1911: 39. [synonymized by Evenhuis & Pape 2017]. Pleciomyia Brunetti, 1911: 269. Type species: Penthetria melanaspis Wiedemann, 1828: 72 (by monotypy). Parapleciomyia Brunetti, 1912: 446. Type species: Parapleciomyia carbonaria Brunetti, 1912: 447 (by monotypy). Nomenclatural notes . The genus Bibiopsis Heer, based on a fossil, has historically been treated as a junior synonym of Penthetria (Loew 1868), as a distinct genus (Scudder 1891), and as a junior synomym of a broadly-defined Penthetria (including Plecia Wiedeman and Protomyia Heer; Handlirsch 1907). More recently, Evenhuis (1994, 2014) treated Bibiopsis as a junior synonym of Plecia . However, Heer’s (1849; Plate XV, Fig. 24b) illustration depicts a stout-bodied fly with R 2+3 elongate and subparallel to R 4+5, rather than short and angled, which indicates that this taxon belongs to the genus Penthetria . Generic Diagnosis . The following combination of characters will distinguish adults of extant members of the genus Penthetria from other Bibionidae: Fore tibia simple (without apical mucron or spines), R 2+3 elongate and subparallel to R 4+5, and antennal flagellomeres short, compact. The larva of Penthetria (Fig. 58) can be distinguished from the larva of other bibionids by the combination of the following characters: presence of fleshy tubercles (each abdominal segment with two transverse rows of two tubercles each), posterior spiracle on segment eight, posterior spiracle with a single ecdysial scar, and the mentum not fused to the anterior margin of the cranium or to the posterior margin of the labium (Fitzgerald 2004, Fig. 36b; pm). Pupae (Figs. 59–60) can be distinguished from all other bibionid genera except Plecia by the presence of fleshy tubercles. Discussion supporting generic diagnosis . The generic diagnosis presented above is based on previous phylogenetic and descriptive work on the family Bibionidae (Fitzgerald 2004) and readers are referred there for further information concerning material examined or the broader context from which the diagnosis was developed. The diagnosis is designed for extant species, but reference to problems and exceptions when diagnosing fossil species are provided below. Additionally, characters not necessarily considered diagnostic, but that may be useful in helping to distinguish the genus, are also discussed. The relatively large adults of Penthetria are usually matte-black, with dark wings. They are sometimes confused with some of the large species of tropical Sciaridae, but are easily distinguished from them by the absence of a narrow eye bridge and the presence of vein R 2+3: in Penthetria males are holoptic and females broadly dichoptic (Fig. 1) (though the odd Palearctic species, P. funebris Meigen, has males with eyes that do not quite meet dorsomedially). Adults of Penthetria are distinguished from other Bibionidae by the apically simple fore tibia (Bibioninae all have apex of fore tibia modified into a long mucron or a circlet of shorter spines), relatively short antennae with compact antennal flagellomeres ( Hesperinus with flagellomeres elongate), and vein R 2+3 elongate and subparallel to R 4+5 (e.g., Figs. 2, 13, 21b, 23) (R 2+3 absent in Bibioninae). The elongate R 2+3 subparallel to R 4+5 is the character typically employed in keys for distinguishing Penthetria from the similar genus Plecia , which has R 2+3 shorter and more oblique or vertical (Fig. 22). Despite the fact that some species/specimens of Penthetria have R 2+3 approaching the condition found in Plecia (e.g., compare Figs. 21a & 22a) and that the form of R 2+3 is essentially evaluated by gestalt rather than any discrete measurement, the extant species can typically be distinguished in this way with confidence. The use of R 2+ 3 in distinguishing these genera becomes more problematic with fossils, where there is a continuum in the form of R 2+3 between typical Penthetria and Plecia (Collomb et al . 2008, and e.g. see discussion of Penthetria by Skartveit & Nel 2017). It may be useful, based on extant taxa, to develop a more objective way of describing the form of R 2+ 3 in order to help more accurately place fossil taxa. For example, in the wing variation of the extant species presented in Figs. 21–22, Penthetria can be defined by the slope of R 2+3 less than 0.43 and Plecia by the slope of R 2+3 greater than 0.74 (slope defined here as rise/run, where rise = measure of imaginary line between points y and z, and where run = measurement of imaginary line between points x and z (see Fig. 22b)). However, when slope is applied to the continuum of the form of R 2+3 found in fossil species, it is not as clear. For example, if slope is measured for the 15 Plecia and 15 Penthetria wings illustrated by Skartveit & Nel (2017), with a line discriminating Plecia with slope>= 0.50, it would correctly assign alone 27 of the 30 wings (i.e., without evaluating any other aspect of morphology). More variation should be studied before determining whether such a definition could be employed, but utilizing a discrete definition of slope and understanding the breadth of variation in the form of R 2+3 (or other veins) of extant species would undoubtably help to objectify the generic placement of fossil taxa. In addition to differences in the shape of R 2+3, Penthetria and Plecia differ in the form of the male terminalia; the male terminalia in Penthetria are very simple, plesiomorphic (e.g., see Fig. 7), relatively homogenous, with a stout, simple, gonostylus articulating in a socket formed from a relatively elongate gonocoxite. The male terminalia in Plecia are much more complex, show great interspecific variability (see e.g., Hardy 1945, Fitzgerald 1998) and tend to have the tubular, gonostyus-socket-portion of the gonocoxite greatly shortened. However, this is less true in fossil specimens. As Skartveit and Nel (2017) point out, the male terminalia of some fossil Plecia are very plesiomorphic and Penthetria -like (e.g., see Collomb et al . 2008). Additionally, Penthetria has subtle parapsidal sutures (longitudinal sutures on the mesonotum corresponding to the position of dorsocentral setae), while they are often present as distinct grooves in Plecia . Females of the two genera can be also be distinguished by Penthetria having two segmented-cerci and Plecia with one-segmented cerci—though some fossil Plecia (e.g., P. clavifemur Skartveit 2009) have been described with two-segmented cerci. It is also noteworthy that Penthetria and the bibionid genus Hesperinus cannot be reliably distinguished based on wing venation alone (Skartveit 2009), which is critical in situations where the fossil specimen is represented only by a preserved wing. In fossils where more than the wing has been preserved, Penthetria has shorter antennae, with more compact flagellomeres and tends to be more stout-bodied, with shorter legs than Hesperinus . Additionally, males of Penthetria are typically holoptic while both sexes of Hesperinus are dichoptic. See also “Intraspecific Variation” (below) with respect to the delimitation of fossil species. Classification and phylogenetic placement . Some studies have placed Penthetria , along with the genus Plecia , in the subfamily Pleciinae of Bibionidae (e.g., Hardy 1981, Blaschke-Berthold 1994) or in the separate family Pleciidae (e.g., Krivosheina 1986) based, in part, on similarities in the larvae of these genera including the “clearly separated frons and clypeus, presence of an additional lobe of the maxillae, and primitive spiracles” (Krivosheina 1969) or musculature of the male terminalia (Ovtshinnikova 1994). More recent morphological phylogenetic studies, however, have not supported these classifications, instead placing Penthetria in its own subfamily (Penthetriinae), which is considered the most basal lineage of bibionids after Hesperinus ( Hesperinus + ( Penthetria + ( Plecia + Bibioninae ))) (Pinto & Amorim 2000, Fitzgerald 2004). This latter hypothesis was not corroborated by the molecular study by Ševčík et al . (2016), which found Penthetria (along with Hesperinus ) to be less basal within the bibionid family tree (Bibioninae + ( Plecia + ( Penthetria + Hesperinus ))). Skartveit & Ansorge (2020) propose placing Hesperinus and Penthetria together in the subfamily Hesperininae based on the Ševčík et al . (2016) study, which finds these two genera to be sister taxa. Further work is needed to reconcile the contradictory hypotheses of these morphological and molecular studies. Characters to support the monophyly of Penthetria are sparse. Fitzgerald (2004) suggested the larval mesothorax with two pseudosegments as a synapomorphy, but so few larvae are known that this character will have to be further vetted as the larvae of more species are discovered. Comments on generic description . The adult and larval generic description provided below is based, in part, on previous phylogenetic and descriptive work on the family Bibionidae (Fitzgerald 2004) and readers are referred there for further discussion concerning homology of structures, material examined, and the broader context from which the description was developed. Examination of all the New World species, as well as a small number of species from the Palearctic and Oriental bioregions were examined. A sclerotized aedeagus is considered absent in Penthetria by Fitzgerald (2004) and it has been presented this way in the generic description (below). Blaschke-Berthold (1994, Fig. 6) suggests that the aedeagus is fused with the parameres in this genus but whether it is absent or indistinguishably fused, a distinct, separate aedeagal sclerite cannot be identified. Sperm transfer in Penthetria is apparently not achieved via an intromittent organ, but probably by spermatophore (Blaschke-Bethold 1994), as is known in some other Bibionidae (Leppla et al . 1975). Larvae are known for only two species of Penthetria . Aspects of the larva of the Palearctic species P. funebris have been illustrated in various papers—mandible (Hennig 1948), habitus and posterior spiracle (Krivosheina 1962), habitus, posterior spiracle, mandible, and labium (Krivosheina & Mamaev 1967), and labium and maxilla (Fitzgerald 2004). The egg, larva, and pupal stages of P. japonica Wiedemann were described and illustrated by Yuan et al . (2015). The larval description below is based on the study of these two species. Generic description . Adult : Small to moderately large flies 4.0-11.0 mm. Head: Male head dorsoventrally compressed; in dorsal view, slightly wider than long, oval-shaped. Head and antennae black to brown. Male with almost entire dorsal surface of head occupied by broadly holoptic compound eye (Fig. 1), except P. funebris , with compound eyes narrowly separated dorsally by longitudinal strip of frons approximately subequal in width to antennal flagellum. Male compound eye strongly divided into larger dorsal region and smaller ventral region; dorsal region usually lighter in color than ventral region and with larger ommatidia. Division between dorsal and ventral regions of compound eye often distinguished by distinct narrow longitudinal step (the dorsal region of the eye is often taller than the ventral region of the eye; where the dorsal and ventral regions meet laterally, a short step, curb, or groove is often visible as in Fig. 12). Division not marked by narrow, longitudinal, shining, sclerotized band; smooth, triangular area devoid of ommatidia also absent. Female head dorsoventrally compressed, slightly wider than long. Female compound eye round or oval in shape, convex, dichoptic (Fig. 1), broadly separated by frons. Female compound eye with inconspicuous, slight longitudinal depression dividing compound eye, but lacking any difference in size of ommatidia or color between dorsal and ventral regions. Compound eye of both sexes bare or clothed with minute, stiff, or very elongate hairs. Ocellar tubercle well developed and prominent in males, moderately to weakly developed in females. Both sexes with three ocelli arranged in small, equilateral triangle. Female with small tubercle or longitudinal ridge just posterior to antennal sockets. Male frons reduced to small, triangular region between anterior margin of compound eye, posterior margin of antennal sockets bearing minute tubercle (or a minute pit in P. funebris ). Sclerotized rostrum not produced, oral margin somewhat ventral in position. Antenna located anteriorly along oral margin, with 7–10 short, stout flagellomeres. Flagellomeres slightly broader than long, except most basal flagellomere, about two times as long as subsequent flagellomere. Pedicel and flagellomeres with subapical ring of short setae and numerous dense, minute, appressed setae. Apical flagellomere slightly more elongate than preceding flagellomere, with rounded point apically. Maxillary palps with five segments; basal segment minute, third segment thickened with dorsal, diagonal sensory pit with dense, minute, specialized setae; apical palpomere slender, more elongate than previous palpomeres. Clypeus broader than long, never elongate. Head entirely sclerotized ventrally. Thorax (Fig. 3): Precoxal-bridge complete. Basisternum present, presternum absent. Dorsum of thorax smooth, matte, often with thick dark pruinosity, usually black, dark brown, anteriorly black and posteriorly orange, or rarely entirely orange, as in P. indica (Brunetti). Parapsidal sutures subtle and not distinct as in some Plecia . Dorsum of thorax with dense, short, stiff to very elongate hairs, laterally, anteriorly, and in dorsocentral rows posteriorly. Thoracic pleura black to dark brown. Males with minute to moderately long hairs on metakatepisternum, and dorsal half of katepisternum. Anepisternum bare or with cluster of hairs at posterior margin. Meron, laterotergite, and mediotergite bare. Female thoracic pleura generally less hairy than male. Legs: Legs black to dark brown. All legs with five tarsomeres, simple tarsal claws, pulvilli and pulvilliform empodia. Anterior coxa short and not reaching ventral edge of katepisternum; reaching about half length of katepisternum in lateral view, usually with elongate hair. Fore femur relatively slender and more elongate; not short and swollen. Fore tibia elongate, slender, apex unmodified, outer edge not developed into a mucron. Fore tibia distally with a single, short spur. Middle legs unremarkable; with two apical tibial spurs. Hind legs with hind femur swollen apically to more slender and only slightly enlarged apically. Hind tibia slender or swollen apically. Inner surface of hind tibia without elongate field of small, round, black, specialized sensilla. Spurs of hind tibia slender, apically acute, subequal in length, or ventral spur slightly more elongate than dorsal spur. In males, hind tarsomeres vary in shape, from slender, elongate, parallel-sided to slightly swollen and sausage-shaped. Female tarsomeres never swollen; slender to robust. Halter black. Wing (Figs. 2, 13): Length, 4.5-12.5 mm, elongate, reaching tip of abdomen, except brachypterous and distinctly shortened in males of P. funebris . Wing brown to blackish fumose. Costal cell often darker than remainder of wing in taxa with fumose wings. Wing color solid or with grade in intensity, but without distinct markings. Pterostigma oval, usually brown to black. Anterior wing veins typically darker than posterior veins. Wing without macrotrichia, with microtrichia. Anal lobe well developed. Costa ends at or just beyond R 4+5 (not shortened or thickened as Enicoscolus Hardy). Subcosta long, complete. Radius without horizontal microstriations. Rs furcate; R 2+3 present, relatively elongate and subparallel to slightly oblique to R 4+5; base R 2+3 often arising at right angle to R 4+5 then sharply bent in direction of wing tip. Base of R 2+3 sometimes with sub-basal stump (appendix) (e.g., P. appendicula Hardy; Fig. 2). R 2+3 branches from R 4+5 from slightly basal to, even with, or distinctly distal to, r-m crossvein. Length of r-m crossvein much shorter than base of Rs. Posterior end of r-m meeting stem of medial fork (M 1+2) or more distally, connecting directly to M 1 (e.g., P. japonica Wiedemann; Hardy & Takahashi 1960, Fig. 2b). Three branches of M present (M 1, M 2, and M 4). Base of M 4 crossvein-like. CuA and CuP present, reaching wing margin, sometimes meeting distally forming closed cell cua. A 1 short, weak; not extending beyond the small fold at the base of anal lobe. Abdomen: Black to dark brown, with short to elongate hairs. Male abdomen elongate, gradually tapered posteriorly; female abdomen much stouter. Male spiracles 1–7 located laterally on abdominal pleura, except spiracle eight, either absent or, if present, a remnant dorsolaterally at membrane between tergites eight and nine (e.g., P. funebris ). Males with tergites and sternites 1–8 unmodified, females with tergites 1–8 and sternites 1–7 unmodified. Male terminalia (Figs. 4–11, 14–20, 24–29, 33–45): Terminalia slightly dorsoflexed, not rotated. Posterior margin of tergite 9 (epandrium) shallowly to deeply (nearly dividing tergite) emarginated (“epandrial cleft”) medially. Epandrial cleft often U-shaped, with lobes at sides of posterior edge of epandrium (epandrial lobes) typically broad, rounded, sometimes more triangular in shape apically. Anterior edge of epandrium sometimes with a shallow to moderate medial emargination. Gonocoxites fused to sternite 9 (hypandrium) into a continuous genital capsule (synsternogonocoxite). Hypandrium distinguishable only by narrow, strap-like thickening ventrally and sometimes a slight medial seam. Posteromedian margin of synsternogonocoxite sometimes with median hump, a median cleft (ventromedian cleft, Fig. 20, vmc), and/or a pair of weak to strong lobes (ventromedian lo : Published as part of Fitzgerald, Scott J., 2021, Penthetria Meigen (Diptera: Bibionidae): Revision of the New World species and world catalog, pp. 451-500 in Zootaxa 4926 (4) on pages 453-460, DOI: 10.11646/zootaxa.4926.4.1, http://zenodo.org/record/4529512 : {"references": ["Meigen, J. W. (1800) Nouvelle classification des mouches a deux ailes (Diptera L.) d'apres un plan tout nouveau. Perronneau, Paris, 40 pp. https: // doi. org / 10.5962 / bhl. title. 119764", "Meigen, J. W. (1803) Versuch einer neuen GattungsEintheilung der europaischen zweiflugligen Insekten. Magazin fur Insecktenkunde, 2, 259 - 281.", "Meigen, J. W. (1804) Klassifikazion und Beschreibung der europaischen zweiflugligen Insekten (Diptera Linn.). Erster Band. Abt. I & II. 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