Lindra neroides ( Herrich-Schäffer, 1869 ) Zhang & Dolibaina & Cong & Shen & Song & Mielke & Casagrande & Mielke & Grishin, 2023

Zhang, Jing, Dolibaina, Diego R., Cong, Qian, Shen, Jinhui, Song, Leina, Mielke, Carlos G. C., Casagrande, Mirna M., Mielke, Olaf H. H. & Grishin, Nick V., 2023, Taxonomic notes on Neotropical Hesperiidae (Lepidoptera), Zootaxa 5271 (1), pp. 91-114 : 99-101

Lindra brasus ackeryi O. Mielke, 1978 is a junior subjective synonym of Lindra neroides ( Herrich-Schäffer, 1869) , new combination, and Lindra neroides huxleyi O. Mielke, 1978 , new combination

The taxonomic identities for a number of Hesperiidae species named by Herrich-Schäffer (1869) have been a challenge to determine due to overly brief description without giving localities for these species and the lack of type specimens that were either lost or are not recognizable. A few identification errors leading to misapplication of these names have been made ( Evans 1955). When reviewing names proposed by Herrich-Schäffer in the context of additional information, such as Godman’s copies of unpublished drawings made by Pl̂tz, we found that Cobalus neroides Herrich-Schäffer, 1869 , currently in Decinea Evans, 1955 (type species Hesperia decinea Hewitson, 1876 ), has been misidentified, an error we set out to correct here. We searched for the type material of C. neroides in MFNB, which incorporates the collection of Herrich-Schäffer in its holdings, but were not able to find any credible syntypes after carefully checking all Hesperiidae drawers on two separate visits. Therefore, we assume that the syntypes of C. neroides are either lost or unrecognizable.

The original description presented in a form of identification key, directly compares C. neroides , female(s), to Cobalus corope ( Herrich-Schäffer, 1869) , a species (currently in Tigasis Godman, 1900 ) with two extant syntypes (male and female). Because C. neroides types were female(s), we focus on the analysis of the female T. corope syntype ( Fig. 5a ). The two species share the following characters, translating the relevant segments from the Herrich-Schäffer key: “Hindwings unmarked, at most … [with] small … dots in cells 3 (-5) … a spot in cell 2 is shifted much further towards outer margin [not just below the discal cell spot].” The differences between T. corope and C. neroides are, for T. corope : “[Discal cell with] white longitudinal spot above the origin of the veins 3, 4, above [this spot] a fine dot; cell 2 with a narrow vertical slash, 3 with larger [spot], 6 and 7 with smaller decreasing dots. Beneath on the hindwing a middle dot and a series [of dots] in cells 2-6. Below ♀ with a pale smudge in the cell 1 b of the forewing”, and for C. neroides female: “[Discal cell with] small pale dot above its inner vein, cell 2 with a lunule, cell 3 with small square, 6 and 7 with small decreasing dots. Beneath FW as in corope . The dots on the forewing are barely recognizable.”

The female shown in Fig. 5a is a true syntype of T. corope because it is from the Herrich-Schäffer collection according to its label, labeled as “corope ♀ ” (possibly in Herrich-Schäffer’s handwriting), matches the original description (except that the forewing spot in cell 3 is not larger than the “slash” in cell 2, could be a mistake in the description) and is curated as a type specimen in MFNB. Therefore, the appearance of this female would reflect what Herrich-Schäffer considered to be similar to C. neroides . Comparative analysis of this specimen and the descriptions suggests that the Hayward’s hypothesis (1949) that C. neroides , now in Decinea , is conspecific with Hesperia lydora (Pl̂tz, 1882) (type locality in Venezuela)—a hypothesis that was adopted by Evans (1955) and forms the basis for the current taxonomic treatment of these two taxa—is unlikely to be correct. Indeed, the relevant parts of the original description of H. lydora (also given as a key) can be translated as: “forewing with only one hyaline point by the apex … in cells 2 and 3 with square hyaline spots. Underside … hindwing with white dots in cells 2 and 3, the latter also visible above” (Pl̂tz 1882a). These characters are apparent in the published Draudt’s illustration (1921–1924) of C. neroides , which is possibly a copy of the original Pl̂tz drawing. On this illustration, the forewing has two small subapical spots, not one; the spot in cell 2 is moon-shaped, not square; the discal cell is with a spot (not mentioned and not shown in the illustration of H. lydora ), albeit small; and the hindwing is without spots on dorsal side.

To shed light on the identity of C. neroides , we turned to other sources. Pl̂tz included species described by Herrich-Schäffer and M̂schler in his keys and illustrated these specimens in his unpublished drawings ( Godman 1907). Although there is no certainty that Pl̂tz did not misidentify these species or used type specimens in his work, many of these agreed well with the original descriptions and extant type specimens. We found this to be true for C. neroides . Godman (1907), who inspected unpublished drawings by Pl̂tz was not able to find specimens that look similar to C. neroides and even questioned whether this species was American. Hence, the drawing t[afel]. 534 was copied among others, now bound in BMNH ( Fig. 5b ). The drawing generally resembles the female syntype of T. corope and mostly agrees with the original description of C. neroides , except that it does not show any subapical dots. Draudt’s (1921 –1924) version of C. neroides ( Fig. 5c ) is similar, especially the ventral side illustration, which looks like a copy of the original Pl̂tz’s drawing, but dorsal side differs in showing a single subapical dot and being a larger specimen (reduced in Fig. 5c , ventral side to scale). Either the dorsal image was drawn from a specimen (and not Pl̂tz’s illustration), or Pl̂tz’s originals had several specimens illustrated, but not all were copied by Godman’s artist(s). Moreover, in his description of C. neroides, Pl ̂tz (1882b) refers to the forewing apex either with a single point or unmarked. It is likely that at least two specimens were involved, unless a single specimen differed in the number of spots on the left and the right sides.

The most unusual feature in both Godman’s and Draudt’s illustrations of C. neroides is the wedge-shaped (per Pl̂tz’, but more like an arrowhead) spot near tornus of ventral forewing. This character is not common in Hesperiidae , and may be present in some Old World species (e.g., Caltoris Swinhoe, 1893 and Pelopidas Walker, 1870 ), possibly causing Godman (1907) to doubt American provenance of this specimen. A specimen that looks very much like this illustration ( Fig. 5d ), especially due to the rarely occurring arrowhead tornal forewing spots, was found in MFNB drawer in the same column and just following a male and female syntypes of Decinea dama ( Herrich-Schäffer, 1869) . This specimen of unknown provenance is not a syntype of D. dama because it does not agree with its original description, but it is a nearly perfect match to illustrations and the original description of C. neroides . It has two subapical dots (albeit very small), and the spot in forewing cell 2 can be called a “moon” by shape, fully agreeing with the original description in both characters. This specimen is from the Sommer collection (bought by Staudinger in 1873), not from the Herrich-Schäffer collection, and therefore we were not able to directly link it with either C. neroides or its description published in 1869. Judging only from its age, this specimen could have been collected prior to 1869 and might have been seen by Herrich-Schäffer, and there is a remote possibility that it might have been a syntype of C. neroides . It is possible that this is one of the specimens illustrated as C. neroides by Pl̂tz. However, if it was the case, we would need to explain the difference in the number of subapical spots: two in this specimen, two in original description of C. neroides , but one or none according to Pl̂tz (who might have overlooked these small spots, nearly absent on ventral side). In any case, this specimen agrees with all the information available about C. neroides , and most likely is that species. This is not the species currently called Decinea neroides , and it does not even belong to Decinea (see below).

We hereby designate the specimen shown in Fig. 5d as the neotype of Cobalus neroides Herrich-Schäffer, 1869 . We believe that there is an exceptional need to designate this neotype, because the name neroides has been misapplied and its current usage does not agree with the information known about this taxon. Our neotype of C. neroides satisfies all requirements set forth by the ICZN Article 75.3, namely: 75.3.1. It is designated to rectify the taxonomic identity of Cobalus neroides Herrich-Schäffer, 1869 , which has been inconsistent with the information available about this species; 75.3.2. The characters for the taxon have been given in its original description in a form of a key by Herrich-Schäffer (1869) and are detailed above; 75.3.3. The neotype specimen bears the following five rectangular white labels: [Hew], [Coll. Sommer], [Coll. | Staudinger], [Dama | H-Sch.], and [DNA sample ID: | NVG-21114E09 | c/o Nick V. Grishin]; 75.3.4. Our search for the syntypes is described above, it was not successful, and for that reason we consider that the specimens composing the type series of C. neroides are either lost or unrecognizable; 75.3.5. As detailed above, the neotype is fully consistent with the original description, and is an excellent match to Godman’s copy of Pl̂tz’s drawing and Draudt’s illustration; 75.3.6. The neotype is from unknown locality, and the type locality was not specified for C. neroides , however, a likely origin of the neotype is in Southeast Brazil, probably in Rio de Janeiro; 75.3.7. The neotype is in the collection of the Museum für Naturkunde, Berlin, Germany (MFNB).

The neotype of C. neroides , has been sequenced and compared by phylogenetic with other Hesperiidae , revealing that it is placed deep within Lindra Evans, 1955 (type species Carystus simulius H. Druce, 1876 ) ( Fig. 4 magenta within red). Notably, due to genetic similarities, it is conspecific with Lindra brasus ackeryi O. Mielke, 1978 (type locality in Brazil: Rio de Janeiro) and Lindra brasus huxleyi O. Mielke, 1978 (type locality in Brazil: Paraná). For instance, the COI barcodes of C. neroides neotype and the holotype and allotype of L. brasus ackeryi are 100% identical and differ from the holotype of L. brasus huxleyi by 0.15% (1bp). With neroides , ackeryi, and huxleyi clustering at the same species level, the genomic analysis ( Fig. 4 ) leaves Lindra brasus (O. Mielke, 1968) (type locality in Brazil: Distrito Federal), as a distinct species, which becomes monotypic. The holotype of L. brasus shows 3.3% (22 bp) COI barcode difference from the neotype of C. neroides . Therefore, we propose Lindra neroides ( Herrich-Schäffer, 1869) , comb. nov., place Lindra brasus ackeryi O. Mielke, 1978 , syn. nov. as its junior subjective synonym, and recognize its subspecies Lindra neroides huxleyi O. Mielke, 1978 , comb. nov. Finally, on the basis of these genetic similarities and known distributions of these taxa, we hypothesize that the type locality of Lindra neroides (neotype of unknown provenance) is in Southeast Brazil, likely in Rio de Janeiro.

Figures

FIGURE 4. Nuclear genome tree of selected Carystina and Hesperiina. The tree is constructed from protein-coding regions. Statistical support values are shown by nodes. For each specimen, its name adopted in this work is given first, and a previously used name is listed in square brackets (if different), supplemented with the DNA sample number, type status, general locality, year of collection (when known), and, for primary type specimens, their repository. Synonyms are given in parentheses preceded by “=”, and in addition by “‡” for unavailable names. The type status refers to this synonym, if the synonym name is provided. Green vertical bar cuts the tree into genera, except Vertica, which is characterized by lower evolutionary rate (leaves are closer to the root compared to most others). Yellow shading denotes genera and its width (horizontal dimension) is proportional to the genetic differentiation within each genus. Taxa discussed in the text are shown in different colors: Carystus elvira (orange), Orthos minka, stat. nov. (green), Orthos orthos (purple), Orthos hyalinus, stat. rest. (olive), Oxynthes (blue, subgenus Oeonus labeled in dark blue), Lindra neroides, comb. nov. (red, neotype labeled in margenta), Lindra brasus (cyan), and Decinea (gray).

FIGURE 5. Lindra neroides and Tigasis corope. a. Tigasis corope (Herrich-Schäffer, 1869) syntype, ♀ [MFNB]. b. Godman’s copy of unpublished Pl̂tz’s illustration t. 534 of L. neroides [BMNH]. c. Draudt’s published illustration of L. neroides, dorsal image rescaled and ventral flipped and rotated to match Godman’s copy (Draudt 1921–1924). d. Neotype of L. neroides, ♀ designated herein, with its labels [MFNB]. e. Paratype of Lindra brasus ackeryi O. Mielke, 1978, ♀ from Brazil: Rio de Janeiro [OM-DZUP]. Dorsal and ventral images are on the left and right from each letter, respectively. All images are to scale, except the labels that are reduced compared to specimens and illustrations and a smaller scale is shown for them. Photograph b (by N.V.G.) is © of the Trustees of the Natural History Museum London and is made available under Creative Commons License 4.0 (https://creativecommons.org/licenses/by/4.0/) and e by Diego Dolibaina.