Noctiliostrebla dubia ( Rudow, 1871 )

Alcantara, Daniel Maximo Correa, Graciolli, Gustavo & Nihei, Silvio S., 2019, Revision of Noctiliostrebla (Diptera: Streblidae), parasites of bulldog bats (Chiroptera: Noctilionidae: Noctilio), Zootaxa 4560 (3), pp. 483-521 : 492-496

Noctiliostrebla dubia ( Rudow, 1871)

( Figs 4 , 15A )

Lipoptena dubia Rudow, 1871: 122 . Lectotype ♂ (here designated, ZMUH). Type locality: Venezuela. Host: Noctilio leporinus [as N. dorsatus Desmarest ]. Other references: Rudow (1872: 407 –408; English translation of Rudow [1871]); Speiser (1902: 159 –160; synonymized Paradyschiria fusca Speiser, 1900a with L. dubia in error).

Lepopteryx megastigma Speiser, 1900a: 54 –55; plate III (fig. 2). Lectotype ♀ (ZMHU). Type locality unknown. Host: Noctilio leporinus [as N. dorsatus ]. Other references: Wenzel (1976: 114; lectotype designation, synonymy proposed).

Aspidoptera megastigma: Speiser (1900b: 154; new combination) ; Speiser (1902: 159); Bequaert (1942: 88; list, comments, misidentification).

Noctiliostrebla megastigma: Wenzel in Wenzel et al. (1966 : 564 –565; new combination); Wenzel (1970: 11; catalog).

Noctiliostrebla dubia: Wenzel in Wenzel et al. (1966 : 563–564; new combination); Wenzel (1970: 11; catalog); Wenzel (1976: 113–115; comments and list of examined material, fig. 45e); Guerrero (1995: 147–148; diagnosis); Guerrero (1997: 11; catalog); Dick et al. (2016: 792; catalog).

Diagnosis. The female of N. dubia differs from those of other species by having two transverse bare areas without setae in the middle of the dorsal connexivum, sternum II deeply emarginate and epiproct with anterolateral setae longer and thicker than posterior setae. The male differs from those of other species by the sternite II tumescent medially, sternite VI twisted on its axis, a curved gonopod after distal macrosetae and lacking a thorn-like membranous projection and a sclerotized thorn on the dorsal margin of the aedeagus.

Redescription. Measurements (mm; n = 8, 5 ♂♂, 3 ♀♀): HFL: ♂ 0.49 (0.47–0.51), ♀ 0.60 (0.58–0.62); SL: ♂ 0.51 (0.50–0.52), ♀ 0.57 (0.56–0.58); TL: ♂ 0.50 (0.49–0.51), ♀ 0.54 (0.53–0.55); WL: ♂ 0.33 (0.30–0.36), ♀ 0.34 (0.33–0.37); WW: ♂ 0.21 (0.20–0.22), ♀ 0.22 (0.20–0.22).

Thorax. Mesepimeron with 1–2 setae on each plate. Wings with 2–4 setae on median vein. Metasternum with metasternal lobe strongly emarginate; distance between coxal condyle III and metasternal lobe apex twice as wide as the width of metasternal lobe emargination.

Female abdomen. Syntergite I+II with 30–38 setae on each plate of lateral lobe; inner lateral margin longer than wide, 2–3 times longer than anterior lobe, and moderately inclined. Dorsal connexivum with spiracle III on abdominal fold; cluster of setae around spiracle III with setae anterior to spiracle III as longer and thicker as setae posterior to spiracle III, with 2–3 setae posterior to spiracle III longer than setae anterior to spiracle III but not twice the length, and with the longest setae shorter than the longest setae on syntergite I+II but longer than half their length; longitudinal sideband not extending beyond outer lateral margin of spiracle, with the longest setae close to inner margin of spiracle and half the length of longest setae on cluster of setae around spiracle III and twice as long as setae on median dorsal connexivum; dense cover of setae from spiracle III to spiracle IV on median dorsal connexivum; one row of setae posterior to spiracle V between two bare areas on median dorsal connexivum, and one row of setae posterior to spiracle VI with 2–3 pairs of setae longer than longest setae on dense cover of setae of median dorsal connexivum. Tergite VII ( Fig. 4B ) with a conspicuous lobe on posterior margin; wider than epiproct but not twice the width, and at most twice as long as epiproct; inner lateral margin as long as lateral margin of epiproct; with 4–5 setae on each plate. Sternite II ( Fig. 4A ) with discal row of 4–8 setae; posterior margin strongly emarginate, and the longest lateral setae twice as long as the shortest median setae. Sternite VII with 7–9 setae on each plate. Epiproct with two pairs of anterior setae, anterolateral pair thicker than and as long as posterior setae, and anteromedial pair 1/2–1/3 the length of anterolateral pair.

Male abdomen. Syntergite I+II ( Fig. 4D ) with 43–52 setae on each plate of lateral lobe; inner lateral margin moderately concave, wider than long, and of similar length as anterior lobe. Ventral connexivum with segmental setae longer and thicker than other ventral setae but not twice the length. Sternite II ( Fig. 4C ) tumescent at middle, as triangular protuberance, with cover of spiniform setae and row of setae on each side of protuberance; with spiniform median setae, thicker and shorter than lateral setae on posterior margin. Hypopygium ( Fig. 4D ) with setae on ventral margin as longer and thicker as setae around dorsal macrosetae of hypopygium but shorter than dorsal macrosetae of hypopygium; sternite VI as longer and wider as cercus, moderately to strongly curved, and distal half strongly twisted on its axis. Genitalia ( Fig. 4E ) with gonopod slightly curved after distal macroseta, distal half moderately tapered and asymmetric at apex, with the distance between the distal setae and the apex of gonopod twice or more than twice the distance between the distal setae and the dorsal margin of gonopod, but less than 2.5 times that distance; dorsal margin of aedeagus without a thorn-like membranous projection and sclerotized thorn.

Distribution. Brazil (Amazonas), Colombia, Venezuela ( Fig. 15A ).

Host. Noctilio leporinus

Type material examined. Lipoptena dubia ( Figs 4 F–G). LECTOTYPE ♂ [on slide] (ZMUH), here designated: Venezuela: “ Holotype ” [red label], “Lipoptena / dubia / Rudow” [white label with blue border], “ Noctilio / dorsatus / Wenezuela” [white label with blue border], “Aspidoptera / megastigma / Speis. / Dr. Speiser / determ. 25.II.1902 ” [front of white label], “Types ♂, ♀ / fide RL Wenzel / “Ectoparasites of Panama ”, p.564 / ♂ terminalia / mounted / separately” [back of white label], “ Noctiliostrebla dubia (Rudow) / ♂, ♀ Syntypen von Lipoptena dubia / Rudow von Noctilio dorsatus / (= N. leporinus ), Venezuela / Rudow 1871, S. 122 / 1872, S. 407 / Speiser 1902, S. 159–160: Paradyschiti / dubia Speiser 1900 , error / Wenzel, Tipton, Kiewlicz 1966, S. 563– / 564: Noctiliostrebla dubia (R.)” [blue cardboard]. PARALECTOTYPE (1♀) [on same slide as lectotype] (ZMUH): same data as lectotype.

Lepopteryx megastigma . LECTOTYPE ♀ [in a vial with glycerin] ( ZMHU): Unknown locality: “Typus” [red label], “ Noctilio / dorsatus” [white label], “Coll. / H.Loew ” [white label], “LECTO-HOLOTYPE / of megastigma / Speis. [=dubia / Rudow] ♀ / T.C. Maa 1962” [white label], “LECTOHOLOTYPE / of megastigma / Speiser (=dubia / Rudow) ♀ = / small female / without head. / R.Wenzel 1976 / ♂ = dubia / Larger ♀ = / Noctiliostrebla / traubi Wenzel ” [white label] . PARALECTOTYPES (1 ♂, 1 ♀) [in same vial as lectotype; female belongs to N. traubi ] ( ZMHU): same data as lectotype .

Additional material examined. Brazil: 1 ♂, Amazonas, Beruri, Reserva de Desenvolvimento Sustentável Igapó–Açu , igapó vegetation along Preto river , 4°40’53.45”S, 61°21’51.54”W, 24.viii.2013, R. Marciente & J.F.T. Andes Jr. leg., on N. leporinus ( MZSP) . Colombia: 6 ♂♂, 4 ♀♀, Meta, El Parque Macarena, junction of Duda and Guayabero rivers, 20.xi.1976, T.O. Lemke leg., on N. leporinus ( FMNH) . Venezuela: 1 ♂, Amazonas, Boca Mavaca, 84km SSE Esmeralda, 7km up Mavaca river , 138m, tropical rainforest, 2.iii.1967, M.D. Tuttle & F.L. Harder leg., on N. leporinus ( FMNH) .

Taxonomic remarks. An excellent review of the problems involving N. dubia can be found in Wenzel et al. (1966) and Wenzel (1976), and does not need to be repeated here. Below we provide only an overview and comments on the contributions that have dealt with the taxonomic status of the name. Rudow (1871) described Lipoptena dubia based on bat flies collected on Noctilio leporinus (as “ N. dorsatus ”) in Venezuela. In his brief description, the author did not designate a holotype and did not mention the number of specimens he had examined. Even though Rudow allocated the species within a hippoboscid genus, he made it clear that the species could have been described in a new genus since it is so different from the other species of Lipoptena Nitzsch. Unaware of Rudow’s (1871) contribution, Speiser (1900a) described Lepopteryx megastigma based on five specimens. He also did not designate a holotype, but his species description was more detailed than Rudow’s. Later, Speiser (1900b) transferred L. megastigma to Aspidoptera. After learning about Rudow’s work, Speiser (1902) examined the syntypes of L. dubia and declared that the series contained three specimens of Aspidoptera megastigma and one specimen of Paradyschiria fusca Speiser, 1900a . Speiser believed that the description of L. dubia was based on the specimen of P. fusca , and wrote “Rudow hat also zunächst das Exemplar von Paradyschiria fusca m. beschrieben, allerdings stellenweise mit Berücksichtigung des andern Stücks, und daun nachträglich eine Bemerkung über dies andere Stück angefügt, welches zu Aspidoptera megastigma m. gehört. Wenn nun auch unter dem Namen Lipoptena dubia zwei verschiedene Species subsumiert sind, muss er doch möglichst der einen erhalten werden, und dies kann nach den vorhergehenden Ausführungen wohl nur Paradyschiria fusca m. sein. Dieser gebührt also nunmehr der Name Paradyschiria dubia Rudow. ” ( Speiser 1902: 160). Because of this, Speiser transferred dubia to the genus Paradyschiria. As part of a revision of Streblidae, Wenzel et al. (1966) examined a slide containing two of Rudow’s type specimens, one male and one female. Since the slide was mounted dry, the authors had to remount it. Both specimens were in poor condition: the abdomen of the female was telescoped and could not be restored, while the male abdomen was damaged during Wenzel’s attempt to dissect it. According to Wenzel et al. (1966), it was clear that Rudow’s description was not based on a specimen of P. fusca . Furthermore, in 1962, T. C. Maa informed Wenzel (see Wenzel et al. 1966: 564) that he had examined Rudow’s and Speiser’s syntypes, and that the species dubia and megastigma were actually synonymous. As Wenzel did not examine the type series of L. megastigma , he erected Noctiliostrebla, transferred dubia and megastigma to the new genus and kept the names as valid until the type series of L. megastigma could be examined. In 1976, Wenzel was able to examine three of the five syntypes of Lepopteryx megastigma , two females and one male. He concluded that one female and the male were clearly representatives of dubia (but see our comment, below, on the identity of these specimens), while the other female belonged to N. traubi . According to Wenzel (1976), Dr. Maa had designated a female identified as N. dubia as the lectotype of L. megastigma by placing a lectotype label with the material. However, the designation was never published. Wenzel (1976) therefore synonymized L. megastigma with L. dubia , and formalized Dr. Maa’s “designation”. During the present revision, we examined the slide containing the two syntypes of L. dubia (images sent by Drs. Kai Schütte and Frank Wieland, ZMUH). In view of the poor condition of the material, we could not see some of the diagnostic characters, but we observed at least three important characteristics that we deem sufficient to identify these syntypes as N. dubia : the deeply emarginate female sternum II, the curvature of the gonopod posterior to the distal macrosetae, and the absence on the dorsal margin of the aedeagus of both a thorn-like membranous projection and a sclerotized thorn. Based on our observations, the male syntype is designated herein as the lectotype of L. dubia . We also examined the type series of L. megastigma , the same three specimens studied by Wenzel (1976). They are in extremely poor condition. The lectotype’s head is missing, its thorax is heavily damaged, and its abdomen is collapsed. It is not possible to observe most of the features, except for the strongly emarginate sternum II. Unfortunately, we cannot be sure of the identity of the lectotype in view of the great similarity between females of N. dubia and N. falsispina sp. n. Only males of these two species can be separated, by examining the aedeagus. The female paralectotype is clearly N. traubi given that it has a moderately emarginate sternite II. The male paralectotype, considered by Wenzel (1976) as belonging to N. dubia , has sternite VI similar to that of N. traubi , but sternite II is not arched and with a pseudoctenidium, as it is in N. traubi . We opted not to dissect the male genitalia due to the poor condition of the specimen. Despite the fact that we have doubts about the identity of L. megastigma , we maintain its synonymy with L. dubia . Regarding the description of L. dubia by Rudow (1871), we agree with Wenzel et al. (1966) that there are no references to Paradyschiria. The reference to wing rudiments can be associated only to Noctiliostrebla, as Paradyschiria is apterous. Rudow’s descriptive section on the episternal and notopleural sutures, also cited by Wenzel et al. (1966), clearly refers to Noctiliostrebla. Therefore, N. dubia is considered valid and megastigma is kept as a junior synonym of dubia .

Other remarks. In the second paragraph of his comments on N. dubia, Wenzel (1976: 113) defined N. dubia and N. traubi as having a “dorsal subapical thorn-like spine on the aedeagus”. However, we think that this comment was made in error, based on the original description and illustration of the genitalia of N. traubi , and on the keys to species provided by Wenzel et al. (1966) and Wenzel (1976). In the description of N. traubi it is clear that the species lacks a dorsal subapical spine on the aedeagus. Noctiliostrebla dubia has been recorded from Bolivia and Peru ( Guerrero 1995; Dick et al. 2007), but we have not examined material of this species from these countries. However, we believe that at least some of the Bolivian specimens previously identified as N. dubia may in fact belong to N. falsispina sp. n., since some areas in Bolivia are close to the distributional range of N. falsispina sp. n. Although Guerrero’s (1995) diagnosis of N. dubia was based on material from Bolivia and Peru, the diagnosis clearly pertains to N. dubia , since the male does not have any structure on the dorsal margin of the aedeagus. However, it is important to add that the main structure used to separate N. dubia from N. falsispina sp. n. is being discussed for the first time in the present work (the thorn-like membranous projection on the dorsal margin of the aedeagus). Even if the material examined by Guerrero (1995) contained specimens of N. falsispina sp. n., it would be difficult to expect that this structure would be included in his diagnosis.

Figures

FIGURE 4. Noctiliostrebla dubia (Rudow), ♀ paralectotype (ZMUH) and lectotype ♂ (ZMUH) (photos kindly taken by Drs Kai Schütte and Frank Wieland, ZMUH). A. Detail of posterior portion of female thorax and abdomen in ventral view, showing metasternal lobe and sternite II. B. Female tergite VII (with setae insertions) and epiproct in dorsal view. C. Detail of male abdomen in dorsal view, showing sternite II. D. Detail of male abdomen in dorsolateral view, showing syntergite I+II and hypopygium. E. Male genitalia in lateral view. F. Slide with both specimens and labels with identifications made by Rudow (labels on the slide) and by Wenzel (label outside of the slide). G. Detail of label with identification made by Speiser. Abbreviations: emar., emargination of sternite II; i.l.m., inner lateral margin; l.l., lateral lobe of syntergite I+II; mts.l., metasternal lobe; sp.I, spiracle I; sp.II, spiracle II; sp.VII, spiracle VII; st.II, sternite II; st.VI, sternite VI.

FIGURE 15. Distributional maps of Noctiliostrebla species. A. N. dubia (Rudow). B. N. traubi Wenzel. C. N. maai Wenzel. D. N. aitkeni Wenzel.