Nothochrysa ehrenbergi Tauber

Tauber, Catherine A., 2019, South American Nothochrysinae (Neuroptera, Chrysopidae): I. Description of Nothochrysa ehrenbergi sp. nov., ZooKeys 866, pp. 1-18 : 4-13

Nothochrysa ehrenbergi Tauber sp. nov.

Type material.

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 / Tauber 2019 " ( Fig. 7f ).

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. 7f . It refers to Suarius flavescens , a species that now is placed in Chrysopodes (Neosuarius) , and with which the new species shares similar coloration and appearance (see Tauber 2010).

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.

Diagnosis.

Subfamily: This specimen exhibits the following diagnostic features of adult Nothochrysinae (cf.: Tjeder 1966, as Dictyochrysinae; Adams 1967; Brooks and Barnard 1990; Makarkin and Archibald 2013; Breitkreuz 2018): (i) wing-coupling mechanism consisting of a large jugal lobe on the forewing (here, folded ventrally; Fig. 1 ) and a frenulum on the hindwing (here, broken off); (ii) base of the forewing without tympanal organ ( Fig. 1 ); (iii) forewing (and hindwing) with stem of the media extending basally, adjacent to the radius and not fused with it ( Fig. 1a, b ; cf. Breitkreuz et al. 2017: 32); (iv) first intramedian cell triangular, with boundaries formed by the MA, the MP, and the crossvein 1ma-mp ("pseudotriangular", sensu Breitkreuz et al. 2017); (v) pseudomedia ill-defined or appearing to merge with inner (not outer) series of gradates ( Fig. 2 ); (vi) pseudocubitus appearing to merge with outer series of gradates ( Fig. 2 ); (vii) forewing with basal subcostal crossvein present ( Fig. 2 ); (viii) second m-cu crossvein stemming from the proximal half of the first intramedian cell ( Fig. 2 ); (ix) each flagellomere having five or six whorls of setae ( Figs 3e , 3f ); and (x) anterodorsal surface of the metascutum displaying small, convex protrusion ( Fig. 4b ; cf. Breitkreuz 2018, Tauber 2019).

Genus placement: The Chilean specimen under study here falls into the genus Nothochrysa on the basis of the following features of its wings ( Figs 1 , 2 ): (i) forewing and hindwing having well developed pseudomedia and pseudocubitus; (ii) forewing and hindwing with two regular series of gradate veins (inner and outer); (iii) intramedian cell of forewing triangular, elongate, occupying approximately half the width between the pseudomedia and pseudocubitus; (iv) RP of forewing with 10 or more branches ( Adams 1967; Makarkin and Archibald 2013; Archibald and Makarkin 2015; Breitkreuz 2018: 200). [Note: Some specimens of N. californica are known to have only eight or nine branches from the RP.]

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 1 , 2 ; cf. Adams 1967; Aspöck et al. 1980: figs 154, 155; Kovanci and Canbulat 2007: fig. 2): (i) the first anal vein is not forked; (ii) the basal subcostal crossvein is slightly distal to the furcation of the radius; (iii) as in most Nothochrysa species, the first intramedian cell is more wedge shaped than truly quadrangular or triangular (i.e., the MA and MP meet basally at a broadly acute angle); and (iv) the third medial cell (directly below im1, Fig. 2a ) is elongate and extends toward the pseudocubitus well beyond the distal edge of first intramedial cell.

Morphological characteristics.

Head ( Fig. 3 ): Width 1.6 mm (including eyes); ratio of head width to eye width = 3.0: 1. Vertex raised, round; surface pitted anteriorly, with few or no setae, lacking prominent posterior fold. Distance between scapes 0.09 mm; distance between tentorial pits 0.36 mm; length of frons (midway between scapes - midway between tentorial pits) 0.33 mm. Frons relatively wide, with broad longitudinal ridge mesally; surface smooth, shiny, slightly rounded below toruli and at insertion of mouthparts; margin above clypeus straight. Clypeus tapering, with rounded sculpturing basally, indented mesally, slightly expanded distally, with distal margin straight to slightly convex; dorsal surface shiny, smooth, sculptured. Labrum about same width as clypeal margin, with small longitudinal ridge mesally; dorsal surface sculptured, shiny; distal margin bilobed, bearing numerous long setae distally. Antenna 9.7-9.8 mm long (~0.5 × length of forewing); scape shorter than wide (0.23 mm long, 0.33 mm wide), lateral margin straight, mesal margin strongly convex, surface with short setae throughout; pedicel 0.17 mm long, 0.13 mm wide, with numerous short setae; flagellum with basal flagellomeres distinct, somewhat elongate (0.12-0.14 mm long, 0.07-0.08 mm wide), midantennal flagellomeres twice as long as broad (0.15 mm long, 0.07 mm wide), basal two flagellomeres with 4-5 partially indistinct whorls of thickset brown setae extending distally, third flagellomere and others distally all with five distinct whorls of thickset, brown setae extending distally, 0.3 –0.5× width of flagellomere, distal whorl with one or two slender, elongate (~0.75 × width of flagellomere), pale setae extending laterally.

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. 4 ): Cervix not visible. Dorsal thoracic surface with pale longitudinal stripe mesally, probably with broad reddish or brownish stripes or coloration laterally. Prothorax broad, 0.9 mm long, 1.5 mm wide, ratio of length to width = 0.63: 1; pronotum well sclerotized, with textured surface, transverse fold mesally, few or no setae. Legs elongate, slender, probably cream, unmarked, lacking prominent tibial spurs. Tarsus with basal three tarsomeres appearing coalesced, bearing spurs, setae intermixed along undersurface; middle three tarsomeres with expanded lateral lobes bearing spurs, setae in irregular rows; distal tarsomere narrow basally, enlarged distally, bearing numerous elongate, slender, dark setae laterally, distally, terminus bearing pair of claws laterally, large pad mesally; claw amber, with basal enlargement, acute slender hook terminally.

Wings ( Figs 1 , 2 , 5 ): Forewing 18.5 mm long, 6.5 mm wide (at widest point); ratio of length to maximum width = 2.9: 1. Membrane clear, lacking markings; microtrichia present below base of every major vein, pale. Trichosors (sensu Makarkin and Archibald 2013: 140-142) absent. Costal area relatively enlarged; tallest costal cell (7th from base of wing) 1.8 mm tall, 2.7 × width of cell, 0.28 × height of wing; costal crossveins simple, six before 1sc-r, twelve after 1sc-r and before stigma, one (very small) after stigma, none within stigma. Sc extending into stigma, fading but not appearing to merge with C or RA; no crossveins in stigma; first sc-r crossvein slightly distal to R f, slightly basal to M f; RA with one very short veinlet extending to wing margin after stigma. Radial area between RA and RP with single row of ten closed cells; tallest cell (3rd from base of wing) 0.6 × as tall as wide. Intramedian cell (im1 = mamp1) prominent, elongate, triangular, formed by MA, crossvein 1ma-mp, and two abscissae of MP, occupying approximately half the space between MA and CuA, with M f broadly acute, long sides (MA, MP) roughly parallel for most of span; crossvein 2m-cu proximal to midpoint of im1. Three medial cells present (mcu, mcua, mpcua), second, third of these elongate, with roughly parallel sides; MP merging into Psc well beyond im1. Two series of gradate veins parallel basally, diverging slightly medially, converging distally. Approximately nine inner gradates in regular, sinuous series, continuing from Psm in zigzag pattern across center of wing; approximately ten outer gradates continuing from Psc in regular, upturned series. RP with nine marginal forks beyond Psc. Cu furcated after m-cu crossvein, with two closed, four open icu cells. CuA with three furcations before meeting MP; CuP furcated below icu2; thus cubital trace having five terminal veinlets (three from CuA, two from CuP). A1, A2, A3 simple, unforked; a1-a2 and a2-a3 crossveins present; distal part of a3 and jugal lobe with dense patch of microtrichia. Jugal lobe large, quadrate, folded beneath third anal cell, without internal vein; margin bearing long, slender setae basally.

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. 5 ): Membrane clear, somewhat glossy. Stigma slightly opaque, without coloration. Costal, subcostal, radial veins brownish; all other longitudinal veins pale with black marks at intersections and (forewing) at bases of setae. Forewing with posterior veinlets extensively marked black; basal inner gradates pale, others becoming increasingly marked black until entirely black distally; outer gradates mostly black. Hindwing with basal inner gradates pale, marked with black at intersections; outer gradates mostly black.

Abdomen (Male, Fig. 6 ; female unknown): Sclerites, integument of pleural region somewhat soft, flexible; tergites, sternites, pleural region covered with setae of uniformly short length; microsetae present, no microtholi. T6: length 0.78 mm, ~1.8 × height; T7: length 0.80 mm, ~1.6 × height; S6: length 0.67 mm, 0.72 × height; S7: length 0.68 mm, ~0.70 × height. Tergites roughly rectangular, edges acute or slightly rounded, ventral margins straight or slightly concave mesally. Spiracles located approximately in center of lateral membrane, roughly circular externally, not enlarged; atria slightly enlarged, rounded, with bifurcated tracheae. Coloration: body somewhat discolored; setae pale. Tergites probably green, without markings; pleuron mostly tan; sternites with green longitudinal stripe dorsally, tan ventrally; callus cerci white.

Male terminalia ( Fig. 7 ): T8 broadly wedge shaped, with dorsal surface slightly rounded, length 0.83 mm, height 0.49 mm, considerably longer than dorsal surfaces of either T9 or ectoproct; lateral margins tapering inward ventrally, ventral margin roughly straight. T9 and ectoproct separate, not fused; callus cerci ovate, protruding basally from posterior margin of ectoproct, 0.18 mm length, 0.10 mm width, with ~30 trichobothria of various lengths. T9 rectangular, with distoventral margin rounded; elongate, lightly sclerotized ventral apodeme along ventral margin, extending proximally to midsection of A8. Ectoproct dome shaped, rounded distally, slightly convex basally, tightly curved ventrally, sloping dorsally; callus cerci situated on lower proximal margin. S8 and S9 partially fused, without internal ridge; S9 more heavily sclerotized than S8, posterior margin slightly more sclerotized than remainder of sternite. S8+9 (lateral view) with proximal margin straight ventrally, becoming broadly rounded dorsally, distal margin short, straight, ventral margin straight; terminal knob extending well beyond edge of S9, with elongate setae on ventral margin; dorsal surface of knob contiguous with heavy recurrent membrane attached to elongate gonarcal membrane. Subanal plate not found.

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.

Biology.

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 Tauber et al. 2014). Nothochrysa larvae generally are considered debris-carriers, but their packets of debris are small, and their morphology is only moderately modified for debris-carrying. In addition, detailed information on aspects of the developmental and reproductive biology of N. californica is available ( Toschi 1965).

For generic-level comparisons, larval descriptions for genera within Nothochrysinae ( Kimochrysa , Pimachrysa , Dictyochrysa , and Hypochrysa ) have been published (see Tauber et al. 2014). Unfortunately, larvae of Asthenochrysa , Leptochrysa , Pamochrysa, and Triplochrysa are not described.

Known distribution.

Currently, this species has only been reported from the type locality, which presumably is the Valle Las Trancas in the region of Ñuble, Chile.

Etymology.

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).

Characteristics shared with Archaeochrysa species

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 ( Adams 1967; Aspöck et al. 1980: figs 154, 155; Makarkin and Archibald 2013: 135, 136). The feature is variable in Archaeochrysa specimens where A1 is visible. It is not forked in two species ( Adams 1967: 237), forked in two species ( Adams 1967: 230, Makarkin and Archibald 2013: 135), and missing from the specimen of the fifth species ( Archibald and Makarkin 2015: 363).

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 ( Adams 1967, Makarkin and Archibald 2013), but not with other known Nothochrysa species.

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 ( Adams 1967, Makarkin and Archibald 2013, Archibald and Makarkin 2015).

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 ( Makarkin and Archibald 2013: 134). In N. ehrenbergi , MA aligns with RP for about one-third the length of the upper margin of the im1 cell, and no crossvein is present ( Figs 1b , 2b ). However, even with this character there appears to be a possible exception. Figure 2 accompanying the original description of Nothochrysa turcica Kovanci and Canbulat shows a short crossvein between RP and MA; confirmation of the accuracy of this drawing is necessary.

Phylogenetic position of Nothochrysa ehrenbergi sp. nov.

Given the above, Archibald and Makarkin’s (2015) discussion of the phylogeny of Archaeochrysa species is worthy of consideration here. Their paper evaluates how the various Archaeochrysa species express three features; each feature has several conditions ranging from presumably plesiomorphic to more derived. Below, the three features are considered, relative to their expression by Nothochrysa species, especially N. ehrenbergi .

(1) The shape of the im1 cell. Archibald and Makarkin (2015) describe two configurations for this character; N. ehrenbergi expresses the second (more advanced) condition in which the sides of the im1 cell are almost parallel for most of their span and converge basally at a relatively steep angle. The extant species of Nothochrysa , including N. ehrenbergi , share this feature with two species of Archaeochrysa .

(2) The position of crossvein 2m-cu. Archibald and Makarkin (2015) list six conditions for this character, each one considered more evolutionarily advanced than the preceding. Nothochrysa ehrenbergi falls into Condition 5, a derived condition in which 2m-cu is located distinctly in the proximal part of im1 (as shown in fig. 2C of Archibald and Makarkin 2015). This character state is typical of at least two Archaeochrysa species, A. creedi (Adams) and A. paranervis (Adams), as well as several other extant genera in Nothochrysinae, including Nothochrysa .

(3) The crossveins of Psc. Archibald and Makarkin (2015: 366) describe and illustrate four character states for this feature; interested readers are referred to the original paper. Suffice it to say here, N. ehrenbergi , as well as three Archaeochrysa species but no other Nothochrysa species, fall into the second of the four conditions. This position is considered plesiomorphic among Nothochrysinae, both fossil and extant ( Archibald and Makarkin 2015).

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.

Figures

Figure 1. Nothochrysa ehrenbergi sp. nov. (Nuble, 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. A 1, A 2, A 3 first, second, third anal veins Cu cubitus Cu f furcation (division) of cubitus Ju jugal lobe M media MA media anterior m-cu media-cubital crossvein R radius RP radius posterior.

Figure 2. Nothochrysa ehrenbergi sp. nov. (Nuble, 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. A 1, A 2, A 3 first, second, third anal veins CuA, CuP anterior, posterior branches of cubitus icu 1, icu 3 first, third intracubital cells ig inner gradate im 1, im 2 first, second intramedian cells Ju jugal lobe MA media anterior MP media posterior mcua, mpcua second and third medial cells M f furcation of media og outer gradate Psc pseudocubitus Psm pseudomedia R f furcation of radius RP radius posterior RP 1 first branch of radius posterior 1 sc-r first crossvein between subcosta and radius 2 m-cu second crossvein between media and cubitus.

Figure 3. Nothochrysa ehrenbergi sp. nov. (Nuble, Chile; Male, CAS): Head and prothorax (a) head, frontal (b) head and prothorax, dorsal (c) head and prothorax, lateral (d, e) base of antennae, dorsal, lateral (f) flagellar segments, mid antenna.

Figure 4. Nothochrysa ehrenbergi sp. nov. (Nuble, Chile; Male, CAS): Habitus (a) antenna, head, and thorax, lateral (b) mesothorax, metathorax, dorsal (c) metatarsus, dorsal (d) metatarsus, ventral (e) mesotarsus, lateral. p raised metascutal protuberance l. e. mesoscutellar lobate expansion.

Figure 5. Nothochrysa ehrenbergi sp. nov. (Nuble, Chile; Male, CAS): Wings, color slightly enhanced to emphasize pattern of vein markings (a) forewing (b) hindwing.

Figure 6. Nothochrysa ehrenbergi sp. nov. (Nuble, Chile; Male, CAS): Abdomen, cleared (a) midsection-terminus, lateral (b) T 8 (distal), T 9, and ectoproct, lateral (c) terminal abdominal segments, lateral (d) terminal abdominal segments, ventral. apo dorsal apodeme extending below T 8 cc callus cerci ect ectoproct k distal knob extending from S 8 + 9 sr spiracle S 4, S 7 fourth, seventh strenites S 8, S 9 partially coalesced eighth and ninth sternites T 7, T 8, T 9 seventh, eighth, ninth tergites.

Figure 7. Nothochrysa ehrenbergi sp. nov. (Nuble, 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.