Shaanxiana langi, Huang, Hao, 2016

Huang, Hao, 2016, Discovery of a fourth species of Shaanxiana (Lepidoptera: Lycaenidae: Theclinae) from China, Zootaxa 4107 (3), pp. 403-412 : 404-411

Shaanxiana langi sp. n.

( Figs. 3–6 , 9 a, 9b, 9d, 9e, 9g ,, 10, 12, 14)

Type material. Holotype ( Figs. 3–4 ): CHINA: ♂, Guangxi province, Laibin City, Jinxiu County, Dayaoshan Mts., 1000m, 21.V.2015, Song-Yun Lang leg., to be deposited in SNUC. Paratype ( Figs. 5–6 ): CHINA: 1♀, same data as holotype, deposited in Hao Huang’s private collection.

Male holotype description. Length of forewing: 17.8 mm. Eye smooth. Antenna similar to that of S. australis Hsu, 2015 , with black club markedly longer than that of S. takashimai and S. pauper Sugiyama, 1994 . Foretarsus 5- segmented. Hindwing with a long black tail tipped with white, and with anal lobe weakly developed.

Wing-pattern. Upperside. Forewing uniformly dark brown and unmarked; ciliae mostly lost, those remaining dark brown mixed with white. Hindwing dark brown; an ill-defined black marginal spot in space 2; a well-marked orange submarginal patch in space 1c, extending a little into spaces 1b and 2 on both sides; anal lobe black; ciliae lost. Underside. Forewing mostly pale yellow and unmarked except for postdiscal, submarginal and marginal areas; a postdiscal series of silvery-white, crescent-shaped spots continuous from space 1b to space 6, basal to a black submarginal band and a whitish marginal band, the latter with two short black dashes in each space; ciliae mostly lost, those remaining in space 1c white, but those in spaces 3 and 5 brown mixed with white. Hindwing similarly marked as forewing except for marginal and submarginal markings in spaces 1b–2; a distinct black marginal spot in space 2 as on upperside; and an orange submarginal patch present in spaces 1b–1c, extending a little into space 2.

Male genitalia ( Figs. 9 d, 9e, 9g, 10, 12). Tegumen in lateral view as in S. australis , more robust than that of S. takashimai and S. pauper . Uncus in dorsal view protruding medially (differing from S. australis ), but largely concealed by very well developed socii. Socii more developed posteriorly than those of S. takashimai and S. pauper , with posterior margin convex in lateral view. Saccus as long as that of S. takashimai , approximately 1/6 times as long as height of ring (tegumen + vinculum). Falces concealed by socius and lateral process of tegumen in lateral view, gradually hooked near apex as in S. australis , not bent medially as in S. takashimai . Valva with no harpe, in lateral view with no distal process. Juxta U-shaped in posterior view. Phallus similar to that of S. australis , with two carinae penis strongly bent upwards beyond dorsal wall of aedeagus; dorsal wall of aedeagus colorless; entrance of vesica (internal sac) oblique, postero-ventrally downward-facing and very close to the zone on ventral surface of aedeagus; vesica when fully everted extending ventroposteriorly, non-bifurcate, bearing two cornuti, one dorsal, one ventral.

Description of female paratype. Length of forewing: 22.6 mm. Sexual dimorphism well marked in size, wing-shape, ground color of forewing upperside and submarginal markings on both wings underside. Wings broader than the male, the dorsum of the forewing and the costa of the hindwing each relatively longer with respect to the respective termen. Forewing upperside with an extensive, non-iridescent pale blue area from wing-base to postdiscal area, leaving only costal and submarginal areas black. Hindwing ground color brownish-grey, markedly paler than in male, extensively dusted with blue scales in the postdiscal and submarginal areas. Both wings underside with black submarginal band broader than in male, centrally dusted with some obscure, dull-orange scales. Hindwing underside with orange patch in spaces 1b–2 more pronounced than in male. Tails markedly longer than those of male.

Female genitalia ( Fig. 14 ). Lodix longer than that of S. takashimai . 8th tergum in lateral view with anterior margin sinuate. Apophysis anterioris absent. Apophysis posterioris nearly as long as 8th tergum and abruptly enlarged at apex. Papilla analis elongate, depressed dorsoventrally. Lamella developed as a ring around entrance of ostium, with lamella postvaginalis interrupted medially. Antrum abnormal, with a dorsal pouch which is pigmented on its inner wall. Ductus bursae absent, replaced by a cervix bursae immediately connected to antrum. Attachment point of ductus seminalis located on dorsal surface of the beginning of cervix bursae. Cephalic end of cervix bursae sclerotized and pigmented. Corpus bursae global. Signa weakly marked as two groups of sclerotized and pigmented teeth on surface of corpus bursae.

Etymology. This new species is named in honor of Dr. Song-Yun Lang, from Chongqing, who collected the type specimens of the new species.

Diagnosis ( Fig. 9 ). This new species is similar to S. australis , but can be distinguished from the latter by the following combination of male characters (the female of S. australis is still unknown):

1 Size is markedly smaller, with a forewing-length of 17.8mm against 22.2mm in S. australis .

2 Hindwing upperside without a clearly defined orange patch just basal to the black marginal spot in space 2.

3 Black submarginal band on both wings underside less marked by orange scales, although the condition of the specimen examined is fresher in the new species than in S. australis .

4 Hindwing underside without a clearly defined orange patch basal to the marginal spot.

5 Uncus in male genitalia well developed and protruding medially in dorsal view, whereas that of S. australis is weakly developed as “a barely recognizable extension with shallow medial depression” ( Hsu, 2015).

6 Saccus in male genitalia markedly shorter than in S. australis , and nearly as long as that of S. takashimai .

7 Valva of male genitalia, when flattened in full-face view, not strongly arched at upper margin as that of S. australis . (As shown in Fig. 9 , the valvae of both S. langi and S. australis are spread without any part folded, thus a comparison between them is convincing. To give a better understanding for the figures published by Hsu (2015), the valva of a male of S. takashimai was flattened in the same condition and compared with the one given by Hsu (2015); the difference between Fig. 9 h and Fig. 9 i indicates that in the figures published by Hsu (2015), some part of the upper marginal area was folded, not fully spread.)

Remarks. Some of the above-mentioned morphological differences might be explained by individual variation within a single species, such as size, details of wing-pattern, and length of saccus and shape of valva in the male genitalia. However, individual variation affecting the form of the uncus is very seldom seen in the Theclini . The difference between the uncus of S. langi and S. australis is therefore considered indicative of separate species status.

The female paratype of S. langi has nearly the same forewing-length as the holotype of S. australis , however, it has a relatively slender body and shorter antenna than the latter. It is possible that S. australis is a more robust species than S. langi , with a longer antenna and stouter body in proportion to its wing size.

The orange scales of the forewing underside submarginal band of the female S. langi paratype, even though this specimen is in markedly fresher condition than the holotype of S. australis , are less apparent than in the latter.

Generic classification. The discovery of S. australis requires some adjustments to the characterization of Shaanxiana given by ( Hsu, 2015): 1) the uncus can be obsolete or bifurcate; and 2) the harpe can be absent. The female of S. langi demonstrates that marked sexual dimorphism can occur in the genus: the female differs sharply from the male in having extensive pale blue areas on upperside of both wings. A careful examination of both male and female genitalia reveals more specific variations as follows: falces can be concealed by socii and tegumen or entirely exposed; the everted vesica (internal sac) of phallus may be branched or not ( Figs. 12–13 ); antrum of female genitalia can be normal ( Fig.15 C, 15K) or specialized into a dorsal pouch ( Fig. 14 I); ductus bursae of female genitalia can be normal ( Fig. 15 C) or absent and replaced by a cervix bursae ( Fig. 14 C); corpus bursae can be simply spherical ( Fig. 14 C) or bearing lateral pouches ( Fig. 15 K).

The four known species of Shaanxiana clearly divide into two groups: S. takashimai and S. pauper have a bifurcate uncus, less developed socii, exposed falces, an elongate valva with a harpe, less developed carinae penis on the aedeagus, bifurcate everted vesica of the phallus, and in the female genitalia, probably a shorter lodix, a more elongate papilla analis, a normal antrum, a ductus bursae, no cervix bursae, lateral pouches on the corpus bursae, and lacking sexual dimorphism affecting the upperside wing-patterns. In contrast, S. australis and S. langi have a non-bifurcate uncus, more developed socii, concealed falces, a stouter valva without a harpe, better developed carinae penis of the aedeagus, non-bifurcate everted vesica of the phallus, and in the female, probably a longer lodix, stouter papilla analis, a specialized antrum, no ductus bursae, a cervix bursae, no lateral pouches on the ductus bursae, and sexually dimorphic upperside wing-patterns.

Arguably S. australis and S. langi represent a genus separate from Shaanxiana , as the genital differences are comparable to those of other genera of Theclini recognised by Koiwaya (2007). However, there seems little doubt that the two species groups of Shaanxiana together constitute a monophyletic group, and cannot be separated in any phylogenetic analysis using morphological characters, as they share the following genital characters besides their striking exophenotypic similarity: 1) carinae penis paired and well developed; 2) entrance of vesica of phallus large and oblique ventrally; 3) cornuti of phallus paired and well marked; 4) papilla analis elongate and depressed dorsoventrally; 5) lamella ring-shaped and interrupted at middle of lamella postvaginalis; 6) attachment point of ductus seminalis close to entrance of ostium; and 7) signum represented by a group of sclerotized and pigmented teeth on surface of corpus bursae.

The author prefers to use the following indicators (‘rules of thumb’) to delimit genera when there are various choices regarding different clades, giving preference to: 1) the clade with a higher Bremer Support value and resampling support value (such as jackknifing or bootstrapping); 2) the clade with more synapomorphies; 3) the clade with more diagnostic characters, regardless of source (external, internal, larval, molecular etc.); 4) finally, historical usage is preferred if the first three indicators do not help. Koiwaya’s (2007) indicators for generic status were evidently based very largely on genital characters, neglecting many potentially important external features in cases such as Thecla Fabricius, 1807 , and Iozephyrus Wang & Fan, 2002, and of Gonerilia Shirôzu & Yamamoto, 1956 , Cordelia Shirôzu & Yamamoto, 1956 , Neogonerilia Koiwaya, 2007, and Pseudogonerilia Koiwaya, 2007, for which generic separations were rejected by Weidenhoffer & Bozano (2007). As there is no sound work on morphological phylogeny for Shaanxiana , Ussuriana Tutt, 1907 , Gonerilia (sensu Weidenhoffer & Bozano, 2007), Coreana Tutt, 1907 , Laeosopis Rambur, 1858 , and Artopoetes Chapman, 1909 , which were supposed to be close to Shaanxiana in the phylogeny by Koiwaya (2007), the above-mentioned Indicator 3 is preferred in this work. The genus Shaanxiana is separable from related genera not only by the seven genital characters listed above but also by various striking external characters, such as the wing-pattern. Since the number of diagnostic characters of the two species groups of Shaanxiana is less than that of the collective genus, it is at best to treat them as species groups within a single genus.

Field observations. The type specimens were found perching and flying around a tree of Mallotus tenuifolius var. tenuifolius (Euphorbiaceae) at 1:00–2:00 pm on 21st May 2015, at an elevation of 1000m. It was very cloudy and the female eventually captured stayed on the tree when it was disturbed by a strike on nearby branches. The two langi were collected along with five adults of Ussuriana michaelis (Oberthür, 1880) , found on the same tree at the same time.

Figures

FIGURE 9. Morphological differences between S. langi and S. australis, the numbers of arrows correspond to those of the characters under the Diagnosis heading in the text (a, b at same scale; c, d at same scale; e – i at same scale); a, b—wing-pattern of S. langi, male; c—dorsum of S. australis in flattened shape (reproduced from Hsu, 2015 with permission by chief editor of Zootaxa); d—dorsum of S. langi in dorsal view of natural shape; e—male genitalia of S. langi in lateral view of natural shape; f—left valva of S. australis in flattened shape (reproduced from Hsu, 2015 with permission by chief editor of Zootaxa); g—left valva of S. langi, holotype in flattened shape; h—right valva of S. takashimai in flattened shape (reproduced from Hsu, 2015 with permission by chief editor of Zootaxa); i—right valva of S. takashimai in flattened shape, taken from specimen shown in Fig. 7.

FIGURES 1 – 8. Relative size of Shaanxiana species at the same scale (scale bar 1 cm). 1 – 2 — Shaanxiana australis, male holotype (reproduced from Hsu, 2015 with permission by chief editor of Zootaxa); 3 – 4 — Shaanxiana langi, male holotype; 5 – 6 — Shaanxiana langi, female paratype; 7 – 8 — Shaanxiana takashimai; 7 — male from Zhouzhi, Shaanxi; 8 — female from Fengxian, Shaanxi. 1, 3, 5 — upperside; 2, 4, 6 — underside; 7 – 8 — upperside (left half) and underside (right half).

FIGURES 14 – 15. Female genitalia at same scale. 14 — S. langi; 15 — S. takashimai. A = female genitalia in lateral view; B = 8 th tergum, lamella and antrum in lateral view; C = female genitalia in lateral view, with 7 th tergum and 7 th sternum removed; D = lamella and antrum in ventral view; E = 8 th tergum and papilla analis in dorsal view; F = 8 th tergum and papilla analis in ventral view, removed from genitalia; G, H = ostium bursae and lamella in posterior view; I, J = lamella and ostium bursae in dorsolateral view, removed from 8 th tergum; K, L = lamella, ostium bursae and corpus bursae in dorsal view, removed from 8 th tergum.

FIGURES 12 – 13. Phallus at same scale. 12 — S. langi. 13 — S. takashimai. Pll = phallus in left lateral view; Pd = phallus in dorsal view; Prl = phallus in right lateral view; Pv = phallus in ventral view; Pvl = phallus in ventrolateral view.