Research Article |
Corresponding author: Catherine A. Tauber ( cat6@cornell.edu ) Academic editor: Atilano Contreras-Ramos
© 2019 Catherine A. Tauber.
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 (2019) South American Nothochrysinae (Neuroptera, Chrysopidae): I. Description of Nothochrysa ehrenbergi sp. nov. ZooKeys 866: 1-18. https://doi.org/10.3897/zookeys.866.35394
|
A new species, Nothochrysa ehrenbergi sp. nov., is described from Chile; it is the first species of Nothochrysa to be reported from the Southern Hemisphere and only the second from the New World. The genus now contains six extant species as well as two species known from late Oligocene and Miocene fossils. An updated catalog of the valid Nothochrysa species is presented, and three nomina dubia are discussed. The inclusion of the new species in Nothochrysa is well supported by morphological features. However, it and other species currently in the genus also share significant features with Archaeochrysa, an older genus of Nothochrysinae which is known only from the Eocene (Ypresian) to the late Oligocene. It therefore appears that N. ehrenbergi is among the least derived Nothochrysa species, and that the separation of Archaeochrysa from Nothochrysa is open to question and further examination.
Archaeochrysa, Chile, fossils, Green lacewing, wing venation
The family Chrysopidae currently consists of three extant subfamilies. Chrysopinae, with approximately 75% of the known chrysopid genera, is by far the largest (N = ~80 genera). The other two subfamilies combined are much smaller (N =14 genera): Apochrysinae with five genera (
Currently, there are records of four extant genera of Nothochrysinae from the New World, three of which are endemic to the region: Asthenochrysa Adams & Penny and Leptochrysa Adams & Penny (one species each) in South America, and Pimachrysa Adams (five species) in North America. The fourth genus, Nothochrysa McLachlan, is widespread throughout the Northern Hemisphere, but only one species is known from the New World (western North America).
During the last few years, several very interesting specimens of Nothochrysinae from the New World were found in museums. Among these specimens is a new species of Nothochrysa, the first from South America and the first from the Southern Hemisphere. The article here describes this new species and discusses its possible relationships with other genera of Nothochrysinae. Also included among the recently discovered New World specimens is the second known example of Leptochrysa prisca Adams & Penny. A separate article redescribes and provides images of this rare monotypic genus (
The genus Nothochrysa has had a tortuous taxonomic history that is well summarized by
Extant species |
californica Banks, 1892 [North America: southwestern Canada, western USA] |
capitata (Fabricius, 1793) [Europe: widespread; northern Africa: Algeria, Tunisia] |
ehrenbergi sp. nov. [South America: Chile] |
fulviceps (Stephens, 1836) [Europe: widespread] |
sinica Yang Chi-kun, 1986 [Asia: China] |
turcica Kovanci & Canbulat, 2007 [Eurasia: Turkey] |
Fossil species |
praeclara Statz, 1936 [Miocene: Germany] |
stampieni Nel & Séméria, 1986 [Oligocene: France] |
Current usage of terms for veins in neuropteran wings is largely based on the classic studies of tracheal pathways by
The names of crossveins are in lowercase, contain a hyphen, and often begin with a number; for example, 1c-sc is the first (basal-most) costal-subcostal crossvein. Cell names are written in lowercase, italicized, and often appended with a number; e.g., csc1 refers to the basal-most cell between C and Sc. For historical and grammatical consistency, I retained the traditional prefix “intra”, rather than “inter” (as proposed by
The terminal traces of the various major veins were estimated by following the marginal branches basally to their origins on major veins (Fig.
To avoid uncertainty, it is also worthwhile to mention the terms that refer to the orientation of the wing: anterior – toward the elongate margin on the upper (costal) edge of the wing; posterior – toward the elongate margin along the lower edge of the wing; basal or proximal – toward the inner edge of the wing attached to the body; apical or distal – toward the far, outer edge of the wing.
The terminology for other body parts follows common usage.
The holotype (a male) is in the California Academy of Sciences (CAS). Its labels read: [1] “CHILE: Nuble [Ñuble] / Las Trancas / 20/25-II-1980 / Luis E. Pena [Peña]”; [2] “Suarius / flavescens / (Blanchard) / det. N. Penny, 1988”; [3] “HOLOTYPE / Nothochrysa / ehrenbergi /
This single specimen was found in the CAS collection among the unidentified chrysopids. A subsequent search of the collection did not yield additional examples. Norm Penny’s ID label remains on the specimen but was not included in Fig.
When discovered, the specimen was discolored, and its wings were loosely folded around its body. One pair of wings was removed for study and is now attached with water-soluble hide glue to a card mounted on the pin below the specimen. The other pair fell off and was reattached to the specimen with hide glue. The abdomen was cleared and dissected; it is preserved in glycerin within a genitalia vial attached to the pin.
Subfamily: This specimen exhibits the following diagnostic features of adult Nothochrysinae (cf.:
Nothochrysa ehrenbergi sp. nov. (Ñuble, Chile; Male, CAS): Venation at base of wings (a) left forewing, (b) left hindwing. Note the absence of a tympanal organ at the base of R in the forewing, the independent origin and trajectory of M along the base of R (arrows pointing downward, both wings), and the alignment of RP and MA in the hindwing. A1, A2, A3 first, second, third anal veins Cu cubitus Cuf furcation (division) of cubitus Ju jugal lobe M media MA media anterior m-cu media-cubital crossvein R radius RP radius posterior.
Nothochrysa ehrenbergi sp. nov. (Ñuble, Chile; Male, CAS): Wings with selected features labeled (a) left forewing, (b) left hindwing. Marginal traces of major veins demarcated; arrow (hindwing) indicates alignment of RP and MA along upper margin of first intramedian cell. A1, A2, A3 first, second, third anal veins CuA, CuP anterior, posterior branches of cubitus icu1, icu3 first, third intracubital cells ig inner gradate im1, im2 first, second intramedian cells Ju jugal lobe MA media anterior MP media posterior mcua, mpcua second and third medial cells Mf furcation of media og outer gradate Psc pseudocubitus Psm pseudomedia Rf furcation of radius RP radius posterior RP1 first branch of radius posterior 1sc-r first crossvein between subcosta and radius 2m-cu second crossvein between media and cubitus.
Genus placement: The Chilean specimen under study here falls into the genus Nothochrysa on the basis of the following features of its wings (Figs
Species placement: Apart from being the only known Nothochrysa species reported from South America, N. ehrenbergi is distinguishable from other species of Nothochrysa on the basis of a number of wing characters (Figs
Head (Fig.
Head coloration: Scape cream, with reddish spot on distolateral tip; pedicel, flagellum cream, unmarked; thickset setae in whorls mostly brown, elongate setae pale. Vertex cream, possibly tinged red laterally; dorsal torulus yellow to cream, apparently unmarked. Frons cream, probably with reddish tinge laterally below torulus; torulus cream, unmarked. Clypeus cream, possibly tinged red laterally; basal, distal margins straight. Genal mark dark red/brown throughout, extending to tentorial pit. Labrum probably cream. Palpomeres probably mostly cream, somewhat darkened distally.
Thorax (Fig.
Wings (Figs
Hindwing: 12.4 mm long, 4.2 mm wide. Costal area not enlarged; at least 15 c-sc crossveins before stigma, none within or after stigma. Radial area containing single row of eleven closed cells between RA and RP. Gradate veins in two roughly parallel series, slightly divergent distally; approximately seven inner gradates beyond Psm; approximately 11 outer gradates beyond Psc. Psc with nine marginal forks. MA aligned with RP for approximately one-third length of im1. CuA with two furcations before meeting MP; CuP undivided; thus, wing margin having three cubital veinlets (two from CuA, one from CuP). A1, A2, A3 simple, unforked; a1-a2 and a2-a3 crossveins present. Jugal lobe without internal vein, basal margin bearing long, slender setae.
Coloration of forewing, hindwing (Fig.
Abdomen (Male, Fig.
Nothochrysa ehrenbergi sp. nov. (Ñuble, Chile; Male, CAS): Abdomen, cleared (a) midsection-terminus, lateral (b) T8 (distal), T9, and ectoproct, lateral (c) terminal abdominal segments, lateral (d) terminal abdominal segments, ventral. apo dorsal apodeme extending below T8 cc callus cerci ect ectoproct k distal knob extending from S8+9 sr spiracle S4, S7 fourth, seventh strenites S8, S9 partially coalesced eighth and ninth sternites T7, T8, T9 seventh, eighth, ninth tergites.
Male terminalia (Fig.
Nothochrysa ehrenbergi sp. nov. (Ñuble, Chile; Male, CAS): Male genitalia, cleared, and specimen labels (Penny’s identification label not included) (a) gonarcal complex, dorsal (b) gonarcal complex, frontal, tilted (c) gonarcal complex, posterior (d) gonarcal complex, lateral (e) hypandrium internum (f) labels. c comes g.a. gonarcal apodeme g.b. gonarcal bridge g.p. gonarcal process gse gonosetae on membranous gonosaccus mu mediuncus.
Gonarcus delicate, slender, broadly arcuate; lateral apodemes slender, quadrate (lateral view), rounded distally, with short, contiguous processes mesally, extending forward. Mediuncus closely attached to dorsal surface of gonarcal arch, flat, recurved into an almost fully circular hood, with two internal sclerotized “rods” extending roughly in parallel from mediuncal base to tip, converging slightly at tip; base of mediuncus quadrate (dorsal view), occupying approximately one-fourth span of gonarcal bridge; terminus of mediuncus with expanded lateral wings, rounded mesal protrusion. Gonosaccus transparent, immediately beneath gonarcal arch and mediuncus, with approximately 32 short setae on distinct setal bases uniformly distributed in two equal patches. Hypandrium internum small, located on delicate membrane extending well below gonosaccus, consisting of paired, curved lateral arms meeting mesally at narrow, rounded apex; comes lightly sclerotized, extending forward beyond apex. Gonapsis, gonocristae absent.
Nothing is known about the biology or larval morphology of this species. The gut of the N. ehrenbergi specimen did not contain noteworthy contents.
Larval descriptions of several Nothochrysa species are available for comparison if N. ehrenbergi larval specimens were to become available (see
For generic-level comparisons, larval descriptions for genera within Nothochrysinae (Kimochrysa, Pimachrysa, Dictyochrysa, and Hypochrysa) have been published (see
Currently, this species has only been reported from the type locality, which presumably is the Valle Las Trancas in the region of Ñuble, Chile.
This species is named in honor of Ronald G. Ehrenberg, Irving M. Ives Professor of Industrial and Labor Relations and Economics at Cornell University, an esteemed and cherished colleague of the author and her late husband (Maurice J. Tauber).
As shown above, N. ehrenbergi shares many features with other extant Nothochrysa species, and its inclusion in the genus is well supported. However, the species also expresses many features that differ from Nothochrysa and that are shared by at least some of the five species in the fossil genus Archaeochrysa. I discuss four below:
First, in the N. ehrenbergi forewing, vein A1 is not forked, whereas it is forked in all other Nothochrysa species (
Second, in N. ehrenbergi the basal sc-r crossvein arises distal to the furcation of the radius and almost directly above the furcation of the media. Both of these character states are shared with the fossil genus Archaeochrysa (
Third, in N. ehrenbergi the distinction between the inner gradate series and the pseudomedia as well as between the outer gradate series and the pseudocubitus is indistinct. Rather, the gradate series and their respective pseudoveins tend to run together more smoothly as a curve, rather than at an angle as in other Nothochrysa species. Again, this feature of N. ehrenbergi is shared most closely with Archaeochrysa species (
Fourth, currently the primary feature used to distinguish between Nothochrysa and Archaeochrysa is the presence or absence of a crossvein between RP and MA in the basal part of the hindwing. The crossvein is present in all known Archaeochrysa species and is reported to be absent from Nothochrysa (
Given the above,
(1) The shape of the im1 cell.
(2) The position of crossvein 2m-cu.
(3) The crossveins of Psc.
On the basis of the above information, it appears that N. ehrenbergi shares a very close phylogenetic relationship with the fossil genus Archaeochrysa. At this point, only one character (the absence of a crossvein between the RP and the MA above the first intramedial cell of the hindwing) supports its exclusion from Archaeochrysa, and this character may have exceptions within Nothochrysa. Indeed, there does not appear to be a synapomorphic character that consistently differentiates Nothochrysa from Archaeochrysa. Thus, given the overall similarity between N. ehrenbergi and the known Archaeochrysa species, I recommend that future studies examine the validity of maintaining the generic separation.
Thanks to Christopher C. Grinter, Collection Manager, and Robert Zuparko, Curatorial Assistant, Department of Entomology, California Academy of Sciences, for facilitating my visits to the collection and for making them enjoyable. Also, I truly appreciate the careful editorial review by Agatha J. Tauber, as well as the thoughtful suggestions provided by the two reviewers and the editor.
My systematic work continues to benefit from earlier support supplied by the National Science Foundation, the NRI-USDA Competitive Grants Program, the National Geographic Society, Western Regional Project W-4185, and Cornell University.
Although
No published information is available that helps identify the generic placement of this species, and apparently the type specimen has not been found (
The original description of this species is relatively detailed for its time, and it includes two illustrations. The type is reported from Mytilene, a city on the island of Lesbos in the North Aegean region (