Dtsch. Entomol. Z. 70 (1) 2023, 159-174 | DOI 10.3897/dez.70.98280 Gnu Se ER BERLIN Resolving inconsistencies between Plétz’s descriptions and presumed type specimens of some Hesperiidae (Lepidoptera) Jing Zhang!*%, Qian Cong! ?, Leina Song! , Jinhui Shen!*, Théo Léger’, Gerardo Lamas?, Olaf H. H. Mielke®, Nick V. Grishin! 1 Department of Biophysics McDermott Center For Human Growth & Development, University of Texas Southwestern Medical Center, 5323 Harry Hines Blvd., Dallas, TX 75390, USA 2 Department of Biochemistry McDermott Center For Human Growth & Development, University of Texas Southwestern Medical Center, 5323 Harry Hines Blvd., Dallas, TX 75390, USA 3 Department of Eugene McDermott Center For Human Growth & Development, University of Texas Southwestern Medical Center, 5323 Harry Hines Blvd., Dallas, TX 75390, USA 4 Museum fiir Naturkunde Berlin, Leibniz-Institut fiir Evolutions- und Biodiversitaetsforschung, Invalidenstr. 43, D-10115 Berlin, Germany 5 Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, Lima, Peru 6 Departamento de Zoologia, Universidade Federal do Paranda, Caixa postal 19020, 81531-980 Curitiba, Parana, Brazil https://zoobank. org/D64F'5500-9339-47C 6-9AAC-C3E995C8A 176 Corresponding author: Nick V. Grishin (grishin@chop.swmed.edu) Academic editor: Wolfram Mey # Received 2 December 2022 # Accepted 3 March 2023 @ Published 3 April 2023 Abstract Comparing specimens curated in MfN as primary types of Hesperiidae names proposed by Carl Pl6tz with the original descriptions and unpublished drawings reveals a number of inconsistencies that we address. Lectotypes are designated for 7e/egonus labriar- is Butler, 1877, Eudamus jalapus Plotz, 1881, and Apaustus interpunctata Plétz, 1884. Neotypes are designated for Netrocoryne seneca Pl6tz, 1882 and Hesperia irma Plétz, 1882. Hesperia ulphila Plétz, 1883 is treated as a nomen dubium. As a result of these designations, the following are junior objective synonyms: Netrocoryne seneca Plétz, 1882 of 7elegonus labriaris Butler, 1877 and Hesperia irma Pl6tz, 1882 of Pamphila irma Méschler, 1879, the latter two names being homonyms. In all these instances, the orig- inal descriptions are satisfied, and, except for 4. interpunctata, the current application of these names is unaffected. Taxonomically, Callimormus diaeses Schaus, 1902, stat. rest. is a valid species, not a synonym of A. interpunctata, which is a junior subjective synonym of Hutocus vetulus (Mabille, 1883). Key Words biodiversity, classification, nomenclature, pseudotype, stability, taxonomy Introduction Primary type specimens serve as bearers of zoological names. They are essential to define each name objective- ly, connecting it to actual specimens that can be studied and compared to others using phenotypic characters or DNA sequences. With the recent advent of genomic se- quencing successfully applied to century-old Hesperiidae type specimens (Cong et al. 2021), it becomes particu- larly important to ascertain that the specimens curated as primary types in collections are indeed the type spec- imens of the names they are considered to be types of. Mistakes can occur in the original descriptions of taxa or during the curation and labelling of type specimens. Some mistakes may be revealed as inconsistencies be- tween the original descriptions and the appearance of Specimens curated as types, including their label data. These errors need to be addressed and corrected to agree Copyright Jing Zhang et al. 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. 160 with the major principles of zoological nomenclature re- corded in the ICZN Code (ICZN [International Commis- sion on Zoological Nomenclature] 1999). The stability of nomenclature that calls for the preservation of the current usage, particularly for the widely known taxa, is an im- portant consideration. However, it needs to be balanced against historical accuracy that calls for applying a name to the taxon it was intended for by the original author. Carl Plotz [1814-1886] named a large number of Hesperiidae species in the course of several years (Pl6tz 1879a—c, 188la, b, 1882a—d, 1883a—d, 1884a-1, 1985, 1886). Many of Pl6tz’s descriptions were given as parts of identification keys and were not detailed enough to differentiate these species from others. However, Plotz also illustrated most of the Hesperiidae he included in his keys. Unfortunately, the original set of these illustrations could not be located. However, copies of some of Pl6tz’s drawings were assembled by Godman for the species he could not confidently recognise and attribute to speci- mens in his collection (Godman 1907). This compilation of the drawings, which are not particularly accurate (it re- mains unclear whether the originals were more accurate), is stored in the library of the Natural History Museum, London, UK (BMNH). At least a second copy of some of these drawings was made because similar drawings, cut into small cards, are pinned among the specimens of corresponding species in the collection of the National Museum of Natural History, Smithsonian Institution, Washington, DC, USA (USNM). Due to sketchy descriptions and (where present) 1l- lustrations and frequently lost or unrecognisable type Specimens, many of the names proposed by Pl6tz are still poorly understood, and their application to species is hypothetical, largely following a comprehensive revi- sion by Evans that still remains the major source of Hes- periidae identifications (Evans 1937, 1943, 1949, 1951, 1952, 1953, 1955). During an attempt to learn more about Pl6tz’s names and the species they were intended for, we stumbled upon a number of inconsistencies in the data. In particular, these involve discrepancies between original descriptions, illustrations, and specimens, including their label data. Here, we discuss and resolve a number of such cases from the perspective of zoological nomenclature. Materials and methods The specimens were inspected and photographed in the following collections: Natural History Museum, Lon- don, UK (BMNH), Zoological Institute and Museum Greifswald (ZIMG), Museum fir Naturkunde, Ber- lin, Germany (MEN), and Zoologische Staatssammlung Munchen, Germany (ZSMC). Historical documents, such as unpublished drawings, were inspected in BMNH and USNM. Photographs of specimens and illustrations were taken with a Nikon D800 camera through a 105 mm macro lens and processed using Photoshop. Genom- ic sequencing and analysis were carried out using our dez.pensoft.net Jing Zhang et al.: Investigations of Plotz's type specimens developed protocols as previously described: general methodology, specifics of working with historical spec- imens, and phylogenetic tree reconstruction are detailed in the following three publications, respectively (L1 et al. 2019; Cong et al. 2021; Zhang et al. 2021). Results and discussion We present the analysis of discrepancies we stumbled upon dealing with the names proposed by Carl Pl6tz. In each case, we suggest a specific solution chosen as a compromise considering all information available to us. However, first, we discuss general considerations re- garding numbers on labels of type specimens cited in original descriptions. Type specimens and specimen numbers A label with a number is commonly placed on a specimen to enable its cataloguing and referencing. Some of such numbers or labels with numbers may be unique to a single specimen; others denote a series where every specimen in the series has a label with the same number. An example of an unusual specimen-unique numbering system would be the one adopted in the past in MfN. Specimens were divided into series (most of the series had only one to four specimens), and each series was assigned a unique number, but apparently, only one label with this number was printed and pinned to a single specimen in the se- ries, which we call a “header” specimen. Other specimens were placed in the collection, arranged in columns, after (below) the specimen with the number, typically bearing no labels on their pins at all. At times, subsequent curators were adding labels to these label-less specimens with the same number as the header specimen (probably to avoid information loss when the collection is rearranged), but these labels usually appeared different from the original printed number labels. Therefore, there may be a special significance to the header specimens in each series, the specimen that actually bears the label with the number. Specimens with such numbers were cited by Pl6tz in his descriptions as “Mus. Berol. n.” (see below). An exam- ple of a number being the same for a series of specimens is the numbering of type specimens in syntypic series in USNM of the names proposed by Schaus and Dyar. For many of these species, there is more than one specimen with the same number labelled as “type”. Not all specimens with the same number are equal Although, unless there is evidence to the contrary—see Dolibaina et al. (2014) for an example involving Cum- bre cumbre (Schaus, 1902)—all specimens with the same number labelled as “type” for taxa named by Schaus Dtsch. Entomol. Z. 70 (1) 2023, 159-174 and Dyar in the USNM collection should be considered syntypes from the perspective of the ICZN Code, not all these specimens were equal in the eyes of Schaus and Dyar themselves. As evidenced by the handwritten labels on these specimens, both Schaus and, to a lesser extent, Dyar, differentiated between the specimen they consid- ered the “type” and other specimens in the series they identified as conspecific with the “type”. Schaus placed handwritten identification labels on the specimens of his new taxa, putting the genus name as the first line, the spe- cies name as the second, and wrote “Schs” in the third. However, for the specimens he considered types, the third line stated “type Schs”. No “type” word was written on the identification labels of other specimens of the syntyp- ic series by Schaus. Sometimes there are two specimens marked by Schaus as types: male and female. The concept at the time was that each sex should have its type. Dyar was less consistent, but many numbered specimens of his taxa did not bear his identification label with the word “type”, although some did (“Type Dyar” in the third line), and sometimes more than one for each sex. Although all these specimens (even those without handwritten identifi- cation labels) have the same “Type No.” labels, it is clear which specimens were preferred as types by Schaus (and to some extent by Dyar), and this fact should be taken into account for lectotype designations. A unique specimen number does not imply holotype In his works, after the new species names he proposed, Plotz regularly wrote the phrase “Mus. Berol. n.” fol- lowed by a number, presumably printed on a label of the specimen Plétz referred to. Although labels with these numbers are unique to a single specimen (1.e., unless a mistake has been made, only one specimen in MfN has a characteristic label with this number), the numbers themselves are generally not unique to specimens with such labels and denote entire series of specimens, usually from one (in which case both the label and the number are unique) to four, as listed in the handwritten catalogue of the collection that needs to be studied in each case. Some- times (see sections below), not all specimens in a series with the same number are from the same locality. Due to these reasons, we suggest that a reference to a single MfN number in Pl6tz’s description should not be taken as the holotype designation. The word “type” or its equiv- alent was not used, and Plotz made it clear that some of his descriptions were based on more than one specimen but provided only one number. For instance, the descrip- tion of Hesperia ulphila Plotz, 1883 detailed both male and female, describing differences between them (Pl6tz 1883b) (see a dedicated section below). However, only a single number was given: “Mus. Berol. n. 5426”. One can only speculate why a single number was referenced and other specimen(s) was (were) not: there should have been at least two specimens involved, one considered by 161 Pl6tz to be a male and the other presumed female. It is e1- ther because other specimen(s) did not bear a number, or were not in the Berlin collections, or their numbers were omitted by mistake. It is equally likely that all these spec- imens were from the same series under the same number. Regardless of the reason, it is possible, and even likely, that in many other cases where a single specimen number is referred to by Plotz, the description was based on more than one specimen, and the assumption of the holotype should be avoided, as suggested in the ICZN Code Rec- ommendation 73F. We note that in a number of cases, Plotz mentioned two collection numbers in descriptions, apparently when he considered that specimens with different numbers, and therefore from different localities, were conspecific. For instance, the original description of Goniurus pilatus lists two numbers: 5068 (3 specimens from “Bahia”, per cata- logue of the collection) and 5069 (1 specimen from “Suri- name”); and Goniurus velinus refers to 5102 (2 specimens from “Bahia’) and 5103 (2 specimens from “Caracas”’) (Plotz 1881b). Plotz did not mention the number of spec- imens for each collection number, and only the collection numbers were listed. We found the number of specimens (2 specimens for each) in the collection catalogue. Neotype designation for Netrocoryne seneca Plotz, 1882 In his description of Netrocoryne seneca (type locality Brazil), Plotz referred to “Mus. Berol. n. 4865” (Plotz 1882d). A specimen with this number (Fig. la—c) was found in the collections of the MfN, but it does not agree with the original description given by Plotz. The last seg- ment of the key referring to NV. seneca is translated as fol- lows: “Forewing in almost all cells with small, disjointed hyaline spots; the across-dash-like [spot] in cell 1 is shift- ed towards the tornus, those [spots] in cells 2, 3 and the discal cell are deeply indented, there is no hyaline spot in the middle at the costal edge. The hindwing is only slightly elongated, with a round hyaline spot in the discal cell and 2 rusty yellow bands against the outer margin, cut by the dark veins; these bands are less complete on the underside. Colouring cloudy olive-brown, palpi grey. ... t. 160. — Mus. Berol. n. 4865. [forewing length] 26 mm. Brazil.” This description is, however, a perfect fit for the unpublished drawing No. 160, copied by Godman’s deci- sion, preserved in the library of BMNH (Fig. 1d—f). Moreover, both the description and the drawing uniquely refer to a characteristically patterned and readily recognisable species known today as Ectomis (Ectomis) labriaris (A. Butler, 1877) (Li et al. 2019) (type locali- ty Brazil: Amazonas, Labrea, Rio Purus), which due to its unique appearance was once placed in a monotypic genus Heronia Mabille & Boullet, 1912 (Mielke 2005). The synonymy between N. seneca and E. labriaris had already been suggested by Draudt (1921-1924) and fol- lowed by all subsequent authors. The presumed “type” dez.pensoft.net 162 Jing Zhang et al.: Investigations of Plotz's type specimens pseudotype of Netrocoryne seneca Plétz, 1882 Nascus b Ectomis labriaris (=Netrocoryne seneca Plotz, 1882) LT labriaris A. Butler, 1877 NT =seneca Plétz, 1882 Ectomis labriaris (A. Butler, 1877) roteas (Cramer, 1780) (60. Mus Ber 4865. Bras ren. | DNAsample ID: | pyaeen sae | | prey ) NVG—15031F11 | a 2 http://coll. min-berlin.de/u/ } | 940b3. | "1996 | c/o Nick V. Grishin | atecp, S4OP% —— tt Cc 160, Nelrocorywe Seneca Platz. \ 160.Mus Ber, 4865. Brasilien. 1 e Nelrocory ne Ssenec a, Plotz. | f Lala Parse, 7 HOLT 8 oT. Cxcterrianrd | Putte Pipe - BMNH(E) 1236025 | j Figure 1. Netrocoryne seneca Pl6tz, 1882. a—c. Pseudotype of NV. seneca with specimen number 4865 in MfN, which is Nascus broteas: a. Dorsal view; b. Ventral view; c. Labels; d—f. Godman’s copy of the unpublished Pl6tz’s drawing of N. seneca identifiable as Ectomis labriaris: d. Enlarged drawing to scale with specimen images (dorsal/ventral, left/right); e. Segment of a page with the drawing; f. Enlarged caption of the drawing; g—i. The lectotype of 7elegonus labriaris Butler, 1877, which is simultaneously the neo- type of N. seneca, both designated 1n this work: g. Dorsal view; h. Ventral view; i. Labels, the round label 1s shown from above (left) and below (right). All images are to scale, except e. Photographs d-i. (by N.V.G.) are Copyright of the Trustees of the Natural His- tory Museum London and are made available under Creative Commons License 4.0 (https://creativecommons.org/licenses/by/4.0/). of N. seneca with the number 4865 on the label is Na- scus broteas (Cramer, 1780) (type locality Suriname), which is patterned differently (e.g., it does not have even a trace of a hyaline spot in the hindwing discal cell), and therefore is not the specimen described and illustrated by Plotz. While we have not investigated the reasons for the inconsistency between the description reinforced by the illustration and the specimen with the number they are supposed to refer to, we conclude that the MfN specimen with the label “4865” is not a syntype of N. seneca. To correct this problem, we searched for possi- ble syntypes of NV. seneca in the collections known to house Pl6étz’s types: MfN, ZSMC and ZIMG. The MfN dez.pensoft.net collection catalogue lists two specimens for No. 4865, but we could not find this second specimen, which might have been the specimen Pl6tz based his illustration and description on. We also failed to find any other candidate specimens and believe that the syntypes of N. seneca are no longer extant. There is an exceptional need to stabilise nomenclature and define N. seneca objectively because the specimen with the number referred to in the original description is not N. seneca, creating a potential for no- menclatural instability if the identity of NV. seneca is de- termined by this specimen. Moreover, the evidence we assembled is unambiguous in revealing the identity of the species that Plétz described as N. seneca, and we confirm Dtsch. Entomol. Z. 70 (1) 2023, 159-174 the subjective synonymy between N. seneca and E. labri- aris that has been in use for nearly a century. Therefore, we hereby designate a single known syntype of E. /abri- aris, female (Fig. 1g—1), as the lectotype of 7elegonus labriaris Butler, 1877 and also as the neotype of Netro- coryne seneca Plotz, 1882, establishing that the latter is a junior objective synonym of the former. This neotype of N. seneca satisfies all requirements set forth by the ICZN Article 75.3. Requirement 75.3.1: it is designated to clarify the taxonomic identity of this taxon, which may be threatened by a specimen of a dif- ferent species curated as a syntype and bearing a label with the number referred to in the original description of N. seneca. Requirement 75.3.2: the characters for the taxon have been given in its original description by Plotz (1882d) and are re-stated above. Requirement 75.3.3: the neotype specimen bears the following labels: || Type || Labria | R. Purus | 1.X.74. | 77-65 | Traill Coll. || Labria R. Purus | 1/10/74. || T. labriaris | Butler Type. || 669 ||. Requirement 75.3.4: Our unsuccessful search for the syntypes of N. seneca is described above, leading us to conclude that they are lost. Requirement 75.3.5: as detailed above, the neotype is consistent with the origi- nal description and additional information (e.g., copies of Plétz drawings) known about this taxon and the cur- rent synonymy of this name. Requirement 75.3.6: the LT (ee ey em Cecropterus jalapus (P\6tz, 1881) Cecropterus jalapus (Pl6tz, 1881) bud CAYVILA S 163 neotype is from Brazil: Amazonas, Labrea, Rio Purus, and the type locality given for NV. seneca in the original description is “Brasilien.” Requirement 75.3.7: the neo- type is in the collection of the Natural History Museum, London, UK (BMNH). Lectotype designation for Eudamus jalapus Plétz, 1881 In his description of Eudamus jalapus (type locality Mex- ico: Veracruz, Xalapa), Plotz referred to “Mus. Berol. n. 4960” (Plotz 1881a). However, the MfN catalogue lists a single specimen from Suriname identified as “Hesperia creteus var. parmenides”’, and in the collection there is a Telegonus sp. from the creteus group with this num- ber. Specimens with numbers close to 4960 are species of Telegonus. However, a specimen curated as a “Type” of E. Jalapus had a number 4970 (Fig. 2a—c). For No. 4970, the MfN catalogue lists a single specimen of “Hesperia sp.” from “Jalappa” collected by Deppe. Turning to the orig- inal description of FE. jalapus, we translate relevant parts of the key as: “Forewing without hyaline spots. Upperside dark brown. Forewing instead of the hyaline [spots] with several dark grey spots; underside of the forewing with dark spots, that [=underside] of the hindwing with 2 dark 13F¢ | ie | No, e/a | teste Ml. Herln DNA sample ID: | NVG—15032A11 c/o Nick V. Grishin | | cy 2 ; S Sie MFNB_Lep Hesperiidae 82831 <5 Zia fay ae (S) ape | Q0% 2 YO oo Yt 5 rae rae Ex cos 2a az & Oo DNA sample ID: | NVG—21114D03 | c/o Nick V. Grishin Pausrus Yuferpunctars pine | Heote- ae as thifig 79-869. agrees Wi _ Isoteinon, pl. 7 53 Vols, ai. of unpublished drawings. °F.A.H.O6. Callimormus Compared w ith 2 vetula, Plotz’s drawing of 2 Mab. clade osso. Godman-Salvin ) - Coll. 1913.—2. e interpunctata Plétz = =Callimormus vetula, Mab. i w2 j ie eee oor | plese. (753), Bahia and Par. j A992. | ; Jn lapunclata: 4 | | tae Fetes K Figure 4. Apaustus interpunctata Plétz, 1884, specimens are separated by white lines, dorsal (left) and ventral (middle) views, en- larged hindwing venter (right), and labels (around) are shown for each specimen: a. Pseudotype, specimen number 5392 in MfN, Ve- hilius vetula (Mabille, 1878); b. Ulustration of “Callimormus vetula” venter from the plate 189h [6] in Draudt (1921-1924); e. Para- lectotype; d. Lectotype; e. Unlabeled specimen in the same column with the types; f. A specimen from Panama, originally from the collection of J. Peter Maassen; current identifications of these specimens are: a, e. Vehilius vetula (Mabille, 1878); b. Callimormus corus E. Bell, 1941; ¢, d. Eutocus vetulus (Mabille, 1883), and f. Callimormus juventus Scudder, 1872; g, h. Specimens in BMNH suggested looking similar to the original Plotz’s drawing No. 753 of A. interpunctata; i. A section of the Hesperiidae drawer in MfN with specimens identified as A. interpunctata (shown in panels a, c—h); j. Notes by Godman (1907) about the original drawing of A. interpunctata; k. An entry from the MfN collection catalogue for the number 5392. All images are to scale, except enlarged views of hindwing and texts. Gray F in e. indicates that the image 1s flipped (left-right inverted, left hindwing shown). Photographs g, h (by Bernard Hermier) are Copyright of the Trustees of the Natural History Museum London and are made available under Creative Commons License 4.0 (https://creativecommons.org/licenses/by/4.0/). dez.pensoft.net 168 Francis Arthur Heron (1 864—1940), who was an assistant curator in the BMNH, found a different specimen that “agrees with fig. . [sic!] pl. 753, Vol. HI. of unpublished drawings.” (Fig. 4g label highlighted in green). This label also reveals that Plétz’s drawings were bound in at least three volumes. The specimen selected by Heron is similar in general appearance to the one chosen by Godman and additionally reinforces the idea about how A. interpunc- tata looked like. However, it is a less precise match to the original description because only 6 dots (unless a streak in cell CuA2-1A+2A 1s counted as a dot) are present on the ventral hindwing: the dot in the cell M,—M, is not ex- pressed. This specimen was later identified by Evans as Eutocus fabulinus (Pl6tz, 1884). Thus, due to superficial similarities, two different workers selected specimens of two different species as possible candidates for how A. interpunctata might have looked. Draudt (1921-1924) reproduced Pl6tz’s drawing, fre- quently losing in quality, as illustrations of species that were not figured in other sources, and he was not able to get specimens of (e.g., Hesperia ulphila Plotz, 1883, Fig. 3j). Likely following Godman, Draudt listed syn- onymy with A. interpunctata and illustrated his concept of “Callimormus vetula” (1.e., Cobalus vetulus Mabille, 1883, not Pamphila vetula Mabille, 1878, which he treat- ed as “Phlebodes vetula’) (Fig. 4b). This illustration de- picts neither Eutocus vetulus nor resembles specimens in BMNH selected to represent A. interpunctata (Fig. 4h, g), but is identifiable as Callimormus corus E. Bell, 1941 (type locality Brazil: Para) due to the pale veins by the forewing costa and the hindwing spots nearly fused into a band. Therefore, Draudt’s illustration is not likely to be a copy of Plotz’s drawing of A. interpunctata and may not offer additional clues about its identity. Then, we turned to the catalogue of historical speci- mens in MfN. This catalogue consists of several hand- written notebooks with handwriting matching that of Carl Heinrich Hopffer (1810-1876), who was the curator of the entomological collections in the MfN until his death. The catalogue lists species and their localities by their collection numbers, one number per line. If there is more than a single locality for one specimen number, the next locality is listed under the previous. The entry for No. 5392 states: 5392 [collection number] \ Interpunctata N. [species name followed by an unknown abbreviation] \ 4 [number of specimens] \ Bahia / Para [locality] \ Sello / Sieber [collectors] (Fig. 4k). It means that four specimens are listed under the number 5392, but only one of these specimens, the “header” specimen, is expected to bear the original printed label “5392” on its pin. The meaning of “N.” following “Interpunctata” is not totally clear. Spec- imen identification labels of similar style and handwrit- ing (1.e., of Hopffer) are found throughout the collection (e.g., Fig. 4a). For the names already published, Hopffer gives the author’s name abbreviation and cites the ref- erence. The names published after his death (1876) by others, e.g., Pl6tz, or unpublished, are followed by this “N.” It 1s not likely to be an abbreviation of a person’s name, but it could stand for the Latin Nobis, 1.e., “us” or dez.pensoft.net Jing Zhang et al.: Investigations of Plotz's type specimens “of us”, the same as “m.” would mean “mihv’. 1.e., “me” or “of me.” Alternatively, and possibly more likely, “N.” could be an abbreviation for the Latin “Nova” or German “Neue”, simply meaning that the name or species was new. The origin of these names with “N.” is also unclear. The names could have been provided to Hopffer by Plotz, who had already prepared the drafts of his manuscripts with names and drawings before 1876, or the other way around, coined by Hopffer himself and simply used by Plotz after Hopfter’s death. The segment of the drawer with the “Typus” spec- imen with the 5392 label, which is a pseudotype of A. interpunctata, is shown in Fig. 41. There are three sim- ilar-sized brown specimens in the same column under it. These specimens did not have any labels at the time they were located in the drawer. It is most likely that these 4 specimens under the header label [ interpunctata | Plotz | are the four specimens listed in the collection catalogue. Therefore at least some of them were available to Plotz for his study. The header specimen (the pseudotype) bears a locality/collector label with the same wording and in the same handwriting as in the catalogue: [ Bahia Sello | Para Sieber ] and the same name label [ Interpunctata | N. | (Fig. 4a green labels). We hypothesise that these three labels: the number, locality, and name, were the original labels, and they refer to all 4 specimens and not just to the pseudotype header specimen. This explains the two lo- calities (and collectors — Friedrich Sello[w] [1789-1831 ] and Friedrich Wilhelm Sieber [1775—1831]) on the label. It is likely that some of these 4 specimens were from one locality, and some were from the other. Other labels were added at some later time, and the label [ interpunct- | tata Pl. | 5392 | type | is in a different handwriting than the original labels, and because it mentioned the word “type” may not have been written by a German (Fig. 4a). There is the fifth specimen within this interpunctata block (Fig. 41). It is smaller than the four, labelled from Panama, comes from J. Peter Maassen collection (hence collected before his death in 1890) and is identified as “Hesperia interpunctata Hopff[er]. ?” It is not a possible syntype of A. interpunctata because it is from Panama, not from Bahia, but we discuss this specimen here be- cause Plotz also attributed the name interpunctata to Hopffer (as Hpf.). However, the identification label might have been written using Pl6tz’s publication, not before it, when Maassen was attempting to identify his specimens, and the question mark indicates that he was unsure of this identification. In any case, it is likely that this specimen was added to the four original specimens at a later date, maybe when the Maassen collection was being incorpo- rated into MfN holdings and was not mentioned in the original collection catalogue. Legs of all five specimens in MfN identified as A. interpunctata were sampled for genomic sequencing, and sample ID labels were added to them (Fig. 3a, c—f). Thus, the three originally unlabeled specimens gained the DNA sample labels (Fig. 31). The genomic datasets of these Specimens were compared with others using phylogenet- ic trees, and a tree constructed from all protein-coding Dtsch. Entomol. Z. 70 (1) 2023, 159-174 Futocus vetulus|18043G11|LT|Panama:ChiriquiJMFNB Eutocus vetulus|19087A04|Colombia:Valle del Cauca|1992 "Eutocus vetulus|OM76.481|Peru:Madre de Dios|2013 ‘_ utocus vetulus (=interpunctata)|21114C05|LT|no data|MFNB Eutocus vetulus (=interpunctata)|21114C04|PLT|no data|MFNB Eutocus quichua|20086E11|Peru:Satipo|2018 Eutocus facilis|19019F01|Panama|1974 Vehilius stictomenes illudens|21014H04|Mexico:Chiapas|1987 Vehilius vetula|19019B03|Guyana|2000 Mehilius vetula [with interpunctata]|18043B09|Brazil| MFNB #5392 Wehilius vetula [with interpunctata]|21114C06|no data|MFNB Vehilius vetula|21047B09|Brazil:Rondonia|1989 e-allimormus juventus|17092C10|Costa Rica|12-SRNP-20224 Callimormus juventus [with interpunctata]|21114D03|Panama|MFNB Figure 5. Mitochondrial genome tree of Eutocus vetulus (red), Vehilius vetula (blue), and relatives. Five specimens in MfN identified as Apaustus interpunctata are labeled in magenta and the lectotype is highlighted in yellow. The lectotype of FL. vetu- lus is labeled in red. regions of the mitochondrial genome (good to illustrate partitioning into species) is shown in Fig. 5. Both by ge- nomic analysis and by phenotype, and using current tax- onomy, 1* (Figs 4a, 5 NVG-18043B09) and 4" (Figs 4e, 5 NVG-21114C06) specimens are Vehilius vetula (Mabille, 1878), 24 (Figs 4c, 5 NVG-21114C04) and 3" (Figs 4d, 5 NVG-21114C05) specimens are Eutocus vetulus (Mabille, 1883), and the 5" specimen (Figs 4f, 5 NVG-21114D03, from Panama) is Callimormus juventus Scudder, 1872 (type locality in Panama). Identification of EF. vetulus is supported by sequencing of its lectotype designed by Mielke and Casagrande (2002), from Panama, also in MfN (Fig. 5 NVG-18043G11). We see that the four original specimens, including the pseudotype, belong to two dis- tinct species in two different genera. These species share a homonymous species epithet, which, curiously, was pro- posed by the same author: Paul Mabille, and caused some confusion, as detailed by Mielke and Casagrande (2002). The locality label for the four specimens belonging to two species gives two distinct states in Brazil: Bahia and Para (Fig. 4a). It is tempting to speculate that two specimens of one species were collected at one locality and two specimens of the other species were collected at the second locality. Because the V. vetula type local- ity is in Para, it is probable that more specimens of this species were collected around the same time at the same locality. Therefore, it seems likely that the two V. vetula specimens (not agreeing with the description of A. inter- punctata) were collected in Para. Hence, the two other specimens, which are E. vetulus (agreeing with the de- scription of A. interpunctata), were probably collected in Bahia. Notably, only Bahia is mentioned as the locality in the original description, supporting our line of thought. Thus, we suggest that the two E. vetulus were collected in Bahia by Sello and not in Para by Sieber. We note that the original Pl6otz’s drawing listed both localities (“Bahia and Para’) per Godman (1907) (not clear if more than one specimen was illustrated or just one with two localities given for it), thus Plotz was aware of both localities at the time he was making the drawings, but chose only one locality in his description. We can only speculate that initially, Plotz might have considered all these four specimens to be conspecific, and those lacking all 7 dots (i.e., V. vetula from Para) were assumed to be worn or variatons. 169 However, when Plétz wrote the description, he probably restricted the concept of A. interpunctata to the specimens with 7 dots (i.e., &. vetulus) and thus to Bahia only. These speculations suggest that the two V- vetula specimens were not syntypes (i1.e., Plétz did not consider them to be A. interpunctata at the time of his publication). A problem arose when the four specimens were still curated under the same original number in the MfN collection (because nobody re-curated them after Plotz’s description). The header specimen was still bearing the label with the number listed in the original description of A. interpunctata, but Plotz probably no longer considered this header specimen of the series that bears that number to be conspecific with his A. interpunctata. After the description was published, curators placed the red “Typus” label, either directly on this header specimen with the number mentioned in the description or as another header label to the series (as the green Bahia/ Para label) that later found its way on the pin of the header specimen when the collection was under curation. We realise that our line of thought above is highly speculative, but the facts are: (1) two E. vetulus speci- mens out of four specimens in “Mus. Berol.” series “n. 5392” (per collection catalogue) fully agree with what is known, published and unpublished, about 4. interpunc- tata. (2) two V. vetula specimens out of four specimens in “Mus. Berol.” series “n. 5392”, including the header specimen, do not agree with the original description and specimens in BMNH selected as the best match to the original drawing of A. interpunctata. For these reasons, the two E. vetulus specimens are syntypes of A. inter- punctata, and we hereby designate the specimen with better developed 7 hindwing dots and thus in better agree- ment with the original description, illustrated in Fig. 4d and currently bearing a single label [ DNA sample ID: | NVG-21114C05 | c/o Nick V. Grishin ] as the lectotype of Apaustus interpunctata P\6tz, 1884 (type locality Brazil: Bahia) to stabilise nomenclature. This specimen lacks an- tennae and, likely due to dermestid damage, the distal part of its abdomen. The second specimen (NVG-21114C04) is a paralectotype. In summary, we support the original synonymy sug- gested by Godman (1907) and, by the identity of its lec- totype, regard Apaustus interpunctata P\6tz, 1884, syn. rev. as a junior subjective synonym of Eutocus vetulus (Mabille, 1883) (has ventral hindwing discal cell pale spot(s)). As a result, Callimormus diaeses Schaus, 1902, stat. rest. (lacks ventral hindwing discal cell pale spots), which is the only available name for the species that Ev- ans (1955) misidentified as Callimormus interpunctata (Plotz, 1884), is reinstated as a species-level taxon. Neotype designation for Hesperia irma Pl6tz, 1882 In his description of Hesperia irma (type locality Co- lombia), Plotz referred to “Mus. Berol. n. 5234” (Plotz 1882a). The specimen in the MfN collection with No. 5234 (Fig. 6a—c) is curated as “Typus” of H. irma, but dez.pensoft.net 170 it shows a number of discrepancies with the original de- scription. The original description was given as a key, rel- evant sections of which are assembled and translated here: “Antenna — almost always — more than 1/2 as long as the forewing. Wings without hyaline spots. Upperside dark brown or black. Forewing below, past the middle, with a pale, basewards sharply and jaggedly delimited smudge. Hindwing at outer margin and anal margin wide viola- ceous-grey. 19. Jrma Pl. Hesp. t. 270. — Mus. Berol. n. 5234. 17 mm. Columbia.” The specimen No. 5234 is sig- nificantly (by more than 1/5) larger than described (20.5 mm, not 17 mm), its antenna is shorter than half of the forewing, the ventral forewing pattern is different from that of Mnaseas derasa derasa (Herrich-Schaffer, 1870), which Pl6tz paired with H. irma by this character (see be- low), the outer margin of the ventral hindwing is actually brown, not violaceous-grey, but a violaceous-grey triangle covers nearly half of the wing in the postdiscal area, before the submarginal area (=Saum), and the specimen is from “Rio” according to its label, not from Colombia, collected by v[on]. Olffers]. There are a number of other specimens in MfN collected by von Olfers, and they are mostly la- belled from “Rio” as well, and none from Colombia. Ignaz Franz Werner Maria von Olfers (1793-1871) must have collected these specimens in Rio de Janeiro when he was there in 1816 as a German diplomat (Wikipedia contrib- utors 2022). This specimen No. 5234 would be identified currently as Mnaseas sirene (Mabille, 1904). Godman did not mention Pl6tz’s drawing No. 270 in his work (Godman 1907) and a copy of this drawing was not made. However, a copy of Cobalus derasa Her- rich-Schaffer, 1870 (which Plotz incorrectly synonymised with Hesperia cassander Fabricius, 1793) drawing No. 269, a species that Plotz paired with H. irma in his key, was available (Fig. 6e). According to Plétz, both C. dera- sa and H. irma share the pattern of the ventral forewing, which is “past the middle with a pale, basewards sharp- ly and jaggedly delimited smudge.” This pattern with a well-delimited jagged basal border is apparent from the copy of the drawing No. 269 and is indeed present in many specimens currently identified as M. derasa (Fig. 6d, f). However, this pattern is missing in the specimen No. 5234. We also note that the antennae of C. derasa, as drawn by Pl6tz, are longer than half of the forewing, agreeing with the description, not shorter than half, as in the specimen No. 5234 (Fig. 6a, b). Then, we asked the question: how would an ideal match to the original description of H. irma look? Search- ing many different collections and butterfly images, we came up with a good option (Fig. 61). Finding this option is a purely theoretical exercise and would not mean that this exact species is conspecific with H. irma because there could be a number of candidate species; some may not even be closely related to each other, only sharing similar appearance. However, we think that it may be use- ful to present various specimens that may fit the original description of H. irma rather well. This specimen, identi- fied as Mnaseas mapirica (E. Bell, 1930) in the collection dez.pensoft.net Jing Zhang et al.: Investigations of Plotz's type specimens of BMNH from an unspecified locality (possibly from SE Brazil or Bolivia), possesses all the characters men- tioned in the description of H. irma. Its ventral forewing has a smudge sharply delimited basad, and the ventral hindwing is broadly violaceous-grey (=veilgrau) along the outer margin and anal margin. Here, we remind that Plotz used the term “veilgrau” loosely because he also described the mostly pale-brown colour of the Cecropter- us (Murgaria) jalapus (Pl6tz, 1881) ventral hindwing as “veilgrau” (see discussion above). Moschler would prob- ably call this colour violaceous-red (veil-roth) (Moschler 1879), a term that Plotz never used, according to our searches. This MZ. mapirica specimen demonstrates that it is possible to find a fit to the original description of H. irma that is nearly ideal and is better than the specimen No. 5234. However, M. mapirica is a southern species not recorded in Colombia, the type locality of H. irma. Next, we looked for a species in Colombia that matched the original description of H. irma. Such a species, widely distributed over both Americas, is apparently Pamphila irma Moschler, 1879 (type locality Colombia), currently in the genus Cynea Evans, 1955 (Fig. 6g—l). No other op- tions revealed themselves. Notably, the ventral hindwing submarginal area of many Cynea irma specimens can be called “veilgrau”’: it is quite similar to the colour in Polyg- onus savigny (Latreille, [1824]), which Plotz described as veilgrau (Pl6tz 1882a). Even the holotype of P. irma Moschler (forewing length 17.5 mm, Fig. 6j-1), also in MfN, is a reasonable match to the description of H. irma Plotz, although the specimen is worn and damaged. For all these reasons, we conclude (1) that the spec- imen No. 5234 in MfN, which is a female of Mnaseas sirene (Mabille, 1904), is not a syntype of H. irma Plotz, and (2) that according to its description, supplemented with unpublished drawings of similar-looking species, H. irma P\otz is most likely conspecific with Pamphila irma Moschler, 1879. Indeed, Hesperia irma is one of many taxa described by both Moschler and Plotz, and it frequently is a challenge to figure out whether the speci- mens used for both descriptions were the same, at least in part. Moschler was sending specimens to Pl6tz for identi- fication (Moschler 1876), and Plotz might have provided his yet unpublished names to Moschler (or the other way around), drawn these specimens, and used them later in his descriptions. The discrepancies between the specimen No. 5234 and the original description of H. irma Plotz, and the agreement of the holotype of P. irma Moschler, 1879 with the original description of H. irma Plotz suggest that in this instance, at least part of the type material may have been shared by Plotz and Moschler, e.g., it is possible that the holotype of P. irma Moschler, 1879 was also a syntype of H. irma Plotz. And the specimen that bears the label with the No. 5234, which is from “Rio” and not from the type locality of both P. irma Moschler, 1879 and H. irma Plotz (“Columbia”), might have been identified as H. irma Plotz later, possibly by subsequent workers who could have switched the labels between specimens, especially if the original specimen No. 5234 was damaged. Dtsch. Entomol. Z. 70 (1) 2023, 159-174 pseudotype of Hesperia irma Plétz, 1882 Mnaseas derasa derasa (=Pamphila gagatina Mabille, 1891) , HT s irma Moschler, 1879 Ze NT =irma Plotz, 1882 ke a. Cynea irma (Moéschler, 1879) Mnaseas derasa derasa (Herrich-Schaffer, 1870) 171 DNA ainaea ID: NVG—18043B10 http://coll.mfn-berlin.de/u/ c/o Nick V. Grishin 44a049 . | Calter Da] i & Mnhnaseas derasa brunnea (O. Mielke, 1972) Mnhaseas mapirica (E. Bell, 1930) = ‘4 arom Sol Bh at ‘| ~ Be ts nak )e tees Coll Méschl. | B.C.A.Lep.Rhc 896, BG Aten ‘ Coll. Gere staudinger| AE OF: ¥ a} a aa) DNA sample ID http://coll. mfn-berlin.de/u/ | NVG-—15035C08 Mata lies | : | (Ope aO| c/o Nick V. Grishin a | Figure 6. Hesperia irma Plotz, 1882. a—c. Pseudotype of Hesperia irma, specimen number 5234 in MfN, which 1s Mnaseas sirene: a. Dorsal view; b. Ventral view; c. Labels; d. A syntype of Pamphila gagatina Mabille, 1891, currently a junior subjective synonym of Mnaseas derasa derasa, venter, Brazil: Rio de Janeiro [MfN]; e. Godman’s copy of the unpublished Plétz’s drawing No. 269 of Cobalus derasa; f. Allotype of Mnaseas derasa brunnea (O. Mielke, 1972), venter; g. Dorsal and h. Ventral views of Cynea irma Venezuela: Aragua, Rancho Grande, 1100 m, 12-Jun-1985, S. S. Nicolay [USNM]; i. Mnaseas mapirica, no locality label, could be from SE Brazil [BMNH]; j-l. the holotype of Pamphila irma Méschler, 1879, which is simultaneously the neotype of Hesperia irma Plétz, 1882 designated in this work: j. Dorsal view; k. Ventral view; I. Labels. All images are to scale. Photographs e. (by N.V.G.) and i. (by Bernard Hermier) are Copyright of the Trustees of the Natural History Museum London and are made available under Creative Commons License 4.0 (https://creativecommons.org/licenses/by/4.0/). To correct these inconsistencies, we searched for syn- types of H. irma in the collections that are known to house Plétz’s types: MfN, ZSMC and ZIMG. Except for P. irma Moschler, 1879 holotype in MfN, which we could not con- vincingly support as a syntype of H. irma Plotz, we failed to find any candidate specimens and believe that the syntypes of H. irma are no longer extant. There is an exceptional need to designate a neotype of H. irma because a specimen curated as a syntype of this name and bearing a label with the number mentioned in the original description is not a syntype: it does not agree with the original description and the type locality given for this taxon, creating a potential for instability of nomenclature leading to unnecessary no- menclatural changes. To stabilise the nomenclature, we hereby designate the holotype of Pamphila irma Moschler, 1879, a male, as the neotype of Hesperia irma Plotz, 1882, making the two names objective synonyms and thus keep- ing the latter as a junior secondary homonym of the former. This neotype is consistent with both the current usage of the name and the original description of this taxon. This neotype of H. irma satisfies all requirements set forth by the ICZN Article 75.3. Requirement 75.3.1: It is dez.pensoft.net 172 designated to clarify the taxonomic identity of this tax- on, which may be threatened by a specimen of Mnaseas sirene (Mabille, 1904) curated as a syntype and bearing a label with the number referred to in the original de- scription of H. irma. Requirement 75.3.2: The characters for the taxon have been given in its original description by Plotz (1882a) and are re-stated above. Requirement 75.3.3: The neotype specimen bears the following 11 labels: || Origin. || A. mer. | Columb. | Sth. 76. || Type | Verhdlg. zool. | bot. Gesellschaft. | 1878. p. 216. | no:21. || Coll. Moschl. || Jrma | Moschl: || P. b. 64: 3. || 896. || Coll. | Staudinger || B-C.A.Lep.Rhop. | Rhinthon | melius, | Geyer. || \QOR code\ http://coll.mfn-berlin.de/u/ | 3226cd || DNA sample ID: | NVG-15035C08 | c/o Nick V. Grishin ||. Requirement 75.3.4: Our unsuccessful search for the syntypes is described above, leading us to conclude that they are lost. Requirement 75.3.5: as detailed above, the neotype is consistent with the original description of this taxon, other information, such as unpublished drawings, and the current synonymy of this name. Requirement 75.3.6: the neotype is from Colombia, and the type locali- ty given for H. irma in the original description is “Colum- bia.” Requirement 75.3.7: the neotype is in the collection of the Museum fiir Naturkunde, Berlin, Germany (MfN). Finally, curious to learn more about the evolution of Pl6tz’s concept of H. irma before the publication of the name, we consulted Plotz’s archive in ZSMC. In his list of Hesperiidae species and specimens known to him, dat- ed 1870, there is an entry “236. ...... Ms.B 5234. / Rio.” where the dots occupy the place where Pl6tz wrote species names (Fig. 7a). E.g., nearby, there is an entry “232. Cas- 242. Jing Zhang et al.: Investigations of Plotz's type specimens sander —F: e.s. 337. Jon VI 24,1. —Don 136. / Derasa —HS: /Sd.Am.”, a species, which was later placed next to H. irma in the published key (Fig. 7a). It seems that in 1870, Plotz knew about the specimen 5234 and that it was from Rio, but he was not able to place a name on it yet. Curiously, a series of specimens under No. 5233 in MfN are M. derasa derasa. Inspecting Pl6tz’s manuscript containing an early ver- sion of his keys, dated 1876, we find derasa and irma next to each other (numbers 277 and 278) (Fig. 7b), just like in the published version (Fig. 7c). And No. 5234 is men- tioned again, but the locality was changed from “Rio” to “Colombia.” The locality for derasa became “Rio” instead of South America as in the 1870 list. The key it- self matches quite closely the published version, howev- er, with one notable difference. The colour of the ventral hindwing marginal areas is described as “dunkelveilgrau’, not simply “veilgrau’”. Dark violet grey is a better descrip- tion of the colour in the neotype of H. irma, which 1s from Colombia. The manuscript was written in 1876, and the neotype was collected in 1876. We suspect that it was around that time when Pl6tz included specimens looking like Pamphila irma Moschler, 1879 (from Colombia) in his concept of H. irma, possibly prioritising them because “Rio” was changed to “Columbia” that remained the type locality in the published version of the key, but “dark” in the colour description was dropped. The number 5234 was retained, but due to all the reasons discussed in this section above, it is not likely that the specimen from Rio was included in Plotz’s final concept of H. irma, and it is not likely that the original drawing, and therefore, the description were based on this specimen. Coss ct0ver Ker Gy, Sonat, Der t I, 496, «