Research Article |
Corresponding author: Catherine A. Tauber ( cat6@cornell.edu ) Academic editor: Atilano Contreras-Ramos
© 2015 Catherine A. Tauber, Francisco Sosa.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Tauber CA, Sosa F (2015) Rediscovery of Nuvol umbrosus Navás (Neuroptera, Chrysopidae, Leucochrysini): redescription and discussion. ZooKeys 519: 141-153. https://doi.org/10.3897/zookeys.519.5996
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The monotypic leucochrysine genus Nuvol was previously known from three specimens of Nuvol umbrosus Navás, collected in the Atlantic Forest region of Brazil. For many years these specimens have been missing, and the genus has remained without a modern description. Here, the species is redescribed based on two newly discovered specimens (females) from the Amazonian region. The female terminalia are relatively simple, except for the subgenitale, which is enlarged, folded into two sections, and heavily sclerotized. Unique aspects of the wing venation and the unusual pattern of banding on the wings support the retention of Nuvol as a valid genus within the Leucochrysini. There are differences between the Amazonian specimens studied here and the earlier descriptions based on specimens from the Atlantic Forest. These differences may indicate the presence of two distinct, geographically separated species within the genus. However, largely because we do not know the sexes of the earlier specimens, we are treating the differences discovered in the two female specimens as expressions of intraspecific variation.
Chrysopinae, Leucochrysini, generic assignment, interspecific variation, intraspecific variation
Leucochrysini (Neuroptera: Chrysopidae) is a large tribe of green lacewings restricted to the New World. Currently, it includes seven genera in varying states of systematic resolution. One of these genera, Nuvol, has received no modern taxonomic treatment: accurate drawings of the only included species, Nuvol umbrosus Navás, are available solely for the wings; its abdominal and genitalic features have not been described (
At the time of its description, the single type specimen of N. umbrosus remained in the Navás collection in Barcelona (
To help determine whether the genus should be retained as a valid entity within the Leucochrysini, the newly discovered Nuvol specimens are described and their features compared, including the taxonomically important genitalia, with those of other leucochrysines. Unfortunately, the generic placement of Nuvol will remain unresolved largely because our specimens are both females, and the male genitalia, as well as the larvae, are unknown.
The tribe Leucochrysini contains the seven genera and two subgenera listed below. Following each genus name are: (i) the number of species currently included in the genus and (ii) references to recent systematic work dealing with the genus.
Berchmansus Navás, 1913 – (two species;
Cacarulla Navás, 1910 – (monotypic;
Gonzaga Navás, 1913 – (eight species;
Leucochrysa McLachlan, 1868
Subgenus Leucochrysa McLachlan, 1868 (41 species;
Subgenus Nodita Navás, 1916 – (150 species;
Neula Navás, 1917 – (monotypic, no specimens known;
Nuvol Navás, 1916 – (monotypic;
Santocellus Tauber & Albuquerque, 2008 – (two species;
The genus Vieira Navás, 1913, which was previously placed in the Leucochrysini (see
Type-species: Nuvol umbrosus Navás, 1916.
Brazil: States of Rio de Janeiro (RJ), São Paulo (SP), Amazonas (AM), Rondônia (RO), as follows. RJ: type specimen reported in original description, Rio de Janeiro, II-1912 (Navás collection, specimen missing); subsequent specimen reported from Prov. Rio de Jan., 20.X.1906, Coll. V. Bönninghausen. M. H., (
Brotéria (Zoológica) 14: 14–35 (description).
The original description states that the type specimen was collected in “Rio de Janeiro, Febrero de 1912” and that it was retained in the Navás collection. The specimen is not reported to be in the Navás collection now (Monserrat 1995).
At some time, a neotype should be designated for this species. However, we are not doing so with either of the two extant specimens because: (1) They are both from the Amazonian region of Brazil, far removed from the type locality – the Atlantic Forest region of Rio de Janeiro. (2) Our specimens are both females; the sexes of the previously described species are unknown. (3) Our specimens both differ significantly from Navás’ illustration and description in aspects of the fore and hind wing venation (discussed below). Thus, until male and female specimens from near the type locality are available for comparison, we consider it prudent to withhold from designating a neotype.
General (Fig.
Wings (Fig.
Nuvol umbrosus Navás wings: a, c, d, e, f Brazil, Rondônia, CAT b Brazil, Amazonas, FS. Note the unforked apical veinlets, markings, radius turning downward at tip, forewing with four intracubital cells. b1 first upper Banksian cell b’1 first lower Banksian cell icu2, icu4 second and fourth intracubital cells im2 second intramedian cell i.g. inner gradate series m3 third median cell ma median arculus (base of first intramedian cell) Psm pseudomedia R radius Rs radial sector rf origin of radial sector rx1 first radial crossvein Sc subcosta.
Unknown.
(Figs
Nuvol umbrosus Navás female abdomen. a, b terminal segments, lateral c callus cerci and trichobothria d ninth tergite and ectoproct, dorsal, divided by deep groove e seventh pleuron and sternite, showing spiracle and texture of integument f callus cerci [a, c, d, e: Brazil, Rondônia, CAT; b, f: Brazil, Amazonas, FS]. a anus c.c. callus cerci g.l. gonapophysis lateralis sg subgenitale sp spermatheca sp.d. spermathecal duct S7 seventh sternite T7 seventh tergite T9+e fused ninth tergite and ectoproct.
Nuvol umbrosus Navás female genitalia. a spermathecal complex b colleterial complex [Brazil, Rondônia, CAT]. b.c. bursa copulatrix b.gl. bursal gland col.d. colleterial duct col.gl. colleterial gland col.res. colleterial reservoir inv invagination sp spermatheca sp.d. spermathecal duct vel velum.
Nuvol umbrosus Navás female genitalia. a, b spermathecal complex and subgenitale, lateral c, d subgenitale, ventral [a, c Brazil, Amazonas, FS b, d Brazil, Rondônia, CAT]. l.l. lower layer of subgenitale sg subgenitale sp spermatheca sp.d. spermathecal duct S7 seventh sternite u.l. upper layer of subgenitale.
Unknown.
Unknown.
Other than differences in color, the two Amazonian specimens examined are very similar to each other, and we are confident that they are conspecific. The coloration of the specimen from Amazonas (INPA) appears deeper than the one from Rondônia (USU) that might be slightly teneral.
Nuvol’s generic relationships. In the original description (
At this time when the anatomy of females from only a small percentage of leucochrysine species has been described, our study does not provide any novel insights into the relationships of this species with other leucochrysines. The internal genitalic structures of the Nuvol female are relatively simple, and they lack the complex, coiled bursal duct of Berchmansus. Thus, they resemble those known for females in a large number of other leucochrysine species. However, the heavy sclerotization and folding of the subgenitale are conspicuous and may be distinctive among leucochrysines. Thus, at this time, there is no compelling evidence that supports a change in the generic designation of Nuvol, and we consider that the external features above provide substantive evidence for a generic difference between Nuvol and other leucochrysines. We await the discovery of Nuvol larvae and broadly based comparative adult morphological and/or molecular studies of the Leucochrysini before making any generic level changes.
Specific identity. It is noteworthy that the two Amazonian specimens we studied differed somewhat from the images and descriptions of the Atlantic Forest specimens studied by
Although it is quite possible that the above venational and color differences are expressions of interspecific variation, at this time we cannot exclude intraspecific, i.e., geographic or sex-associated variation. Our two Amazonian specimens are females; whereas the sexes of the specimens from the Atlantic Forest are unknown. And we note that leucochrysines are notorious for polymorphisms in body color and markings (
In the most recent taxonomic key for chrysopid genera (
82 | Ectoprocts not fused dorsally, or with deep dorsal groove (Fig. 22; also see Fig. |
82A |
– | Ectoprocts fused dorsally; forewing usually unmarked; longitudinal veins forked apically | 84 |
82A | Ectoprocts separated by dorsal suture; forewing with black spots on dcc and pterostigma (Fig. 280); claws with basal dilation (Fig. 11); basal lobe of subgenitale with V-shaped indentation (Fig. 286) | Chrysemosa Brooks & Barnard |
– | Ectoprocts separated by deep dorsal groove; forewing unmarked or marked with numerous small black spots or four large brown bands throughout the wing (Figs 456, 519); claws with or without basal dilation; basal lobe of subgenitale without V-shaped indentation | 83 |
83 | Forewing unmarked or marked with numerous small black spots throughout wing (Fig. 456); claws undilated (Fig. 12); basal lobe of subgenitale elongate (Fig. 460) | Suarius Navás |
– | Forewing marked extensively with large brown bands (Fig. 519; also see Fig. |
Nuvol Navás |
We are indebted to Maurice J. Tauber (Cornell University, deceased) for his encouragement and help during the initial phases of this study. Thanks to Marcio Luiz de Oliveira, INPA Curator of Invertebrates, and the “Fundação de Amparo à Pesquisa do estado do Amazonas” (FAPEAM) Edital: N. 011/2013 - COLEÇÕES BIOLÓGICAS, Protocol: 16066.UNI423.6372.29052013 for hosting the visit of FS to the INPA. Thanks to Carol Von-Dohlen, Utah State University, for loaning CAT chrysopid specimens and her patience in awaiting their return. The “Lacewing Digital Library” website, developed by J. D. Oswald (Texas A & M University, College Station), was very useful during this study. CAT’s work has benefitted from funding by the National Science Foundation (DEB 0542373, with MJT), USDA/NRI Competitive Grants Programs (9802447, with MJT), Cornell University, and the Cornell University Library.