An illustrated catalogue of the type specimens of Lepidoptera (Insecta) housed in the Zoological Museum Hamburg (ZMH): Part I. superfamilies Hepialoidea, Cossoidea, and Zygaenoidea

Zoological collections remain the main archive of animal biodiversity on Earth, and especially in times of large-scale declines of many groups represent important resources for biodiversity and conservation research. The most important individuals of these collections are the type specimens, which represent the original and unique reference for a species. While a full digitization of most museum collections currently remains utopic, lists of types are an essential resource for researchers to perform taxonomic revisions. Here, we provide an updated type catalogue of the lepidopteran superfamilies Hepialoidea, Cossoidea and Zygaenoidea deposited in the Zoological Museum of Hamburg (ZMH). In this paper, we report 259 specimens labelled as “types” belonging to 36 infrasubspecific taxa (invalid under the ICZN codes), 34 subspecies (16 of valid status) and 13 species (all of valid status). We present a full bibliography of the original descriptions and illustrations for all taxa, aiming to provide a comprehensive taxonomic guide to this collection.


Introduction
Natural history collections are the main archives of biodiversity. They focus on collecting, maintaining and documenting natural specimens, a crucial task in times of biodiversity decline. The Zoological Museum of Hamburg (ZMH) holds large collections across all animal groups. With approximately five million specimens and several thousand primary types, the entomological collection is among the most important natural history archives in Germany.
The origins of the current collection of the Zoological Museum Hamburg (ZMH) can be traced back to the first half of the 19 th century, as a result from the initiative and interest in natural history of local civilians, merchandisers, traders, and owners of seagoing vessels (Glaubrecht 2018). However, much of the collections of holometabolous insects were destroyed by the incendiary bombing of Hamburg in 1943 (Weidner 1979). Thus, as all documents such as entry books, counting lists and card catalogs were lost and the former magnitude of the collection remains difficult to describe. However, much of the earlier new entries after WW2 are described in Weidner (1974), with the largest and most important collections which have been obtained being those of Georg Warnecke (1943and 1963), Richard Jänig (1949), Fritz Diehl (1950-1972), Hermann Rödinger (1952, the Jungius Gesellschaft (1956), Hanan Bytinski-Salz (1960), Eduard Feldtmann (1960), Günter Albers (1972, the Altona Museum (1965,1973,1976), and Mathilde Linz (1974). More recently the largest newly obtained collections were those of Hans-Jürgen Kelm (2003), Walter Baltruweit (2013), Jörg Roloff (2014), Thomas Tischler (2017), Bernd Heinze (2020), and Karl-Heinz Müller-Köllges (2020). Besides these large collections, many medium and small collections were obtained by the museum. Hence, despite the considerable loss in WW2, presently the ZMH hosts ca. 5 million insect specimens, being placed among the five largest entomological collections in Germany. About 1 million specimens represent the Lepidoptera. However, despite their importance, the entomological collections in Hamburg are not well documented, nor digitized.
Herbert Weidner, a former curator of the collection had published an important series of papers from the 60s until the late 70s, where he documented the collections in much detail (first catalogue -Weidner 1962, last catalogue -Weidner 1979, and the Lepidoptera catalogue Weidner 1974). However, since the work of Weidner more than 40 years have passed without further documentation. Further, Weidner's catalogues were all published in German, hence, are difficult to access for international researchers and lack family group classification, making it difficult to work with them. Therefore, a comprehensive effort to revise the type catalogues has been initiated to give an update on the type material housed in the ZMH in English language (Sartori et al. 2016;Husemann 2018a, b, Harms andDupérré 2018;Monod et al. 2019;Henningsen et al. 2020;Simoes et al. in press).
To make the Lepidoptera collection more accessible to researchers, we performed a detailed inventory initiated by Hossein Rajaei in 2013. As a part of this inventory, the whole collection was thoroughly searched and types were moved to a separate, specially protected type collection. All type specimens were compared with the original descriptions, photographed, databased and relabeled, if required. The original high-quality photographs are available online through the FUNDus! Portal of the University of Hamburg (https://www.fundus. uni-hamburg.de/). The complete data of the type material is provided as a table (Suppl. material 1: Table S1) in this catalogue.

Material and methods
Photography. All images were taken with a digital imaging system with stacking capabilities. The images where either taken with a Canon EOS 5DSR with a 65 mm or 100 mm lens and stacked with Zerene Stacker (PMax algorithm) or with a Canon EOS 6D with a 55 mm or 65 mm lens and stacked with Helicon Focus 5.3.
Taxonomy. All taxonomic changes were made after studying the relevant original description of the type material. Synonyms that have already been proposed are listed and the type material was examined. Classification used herein follows van Nieukerken et al. (2011). The taxonomy of the family Zygaenidae is following the last revisions of the family (Hofmann and Tremewan 1996;Efetov 2001;Efetov andTarmann 1995, 2017). For a full list of synonyms, please refer to the above-mentioned catalogues.
Type and original specimen material. Type and original specimen localities are cited in their original spelling or with additional remarks (in square brackets), if required. All data are transcribed verbatim as they appeared on the respective labels. For unreadable data aquestion mark (?) is used. Double slash (//) separates data on different labels, a single slash (/) separates lines within each label, and semi-colon (;) separates different specimens. Additional comments are placed in square brackets.
The ICZN (1999) terminologies (e.g., holotype, syntype, lectotype, neotype, paratype, paralectotype, allotype, cotype and etc.) are available at the ICZN website (http://iczn.org/content/Glossary) and articles cited in the text can be found below: Infrasubspecific taxon -a taxon at lower rank than that of subspecies. The names of such taxa are not regulated by the Code. Article 45.6.1. -A name is infrasubspecific if its author expressly gave it infrasubspecific rank, or if the content of the work unambiguously reveals that the name was proposed for an infrasubspecific entity; Article 45.6.2. -A name is deemed to be infrasubspecific if its author used one of the terms "aberration", "ab." or "morph". Article 45.6.3. -A name is deemed to be infrasubspecific if it was first published after 1960 and the author expressly used one of the terms "variety" or "form" (including use of the terms "var.", "forma", "v." and "f."); Article 45.6.4. -A name is subspecific if first published before 1961 and its author expressly used one of the terms "variety" or "form" (including use of the terms "var.", "forma", "v." and "f."), unless its author also expressly gave it infrasubspecific rank, or the content of the work unambiguously reveals that the name was proposed for an infrasubspecific entity, in which case it is infrasubspecific; Article 72.7. -Name-bearing types of nominal species-group taxa denoted by new replacement names (nomina nova). If an author proposes a new spe-cies-group name expressly as a replacement (a nomen novum) for an earlier available one, then the two names are objective synonyms; both the nominal taxa they denote have the same name-bearing type despite any simultaneous restriction or application of the new replacement name (nomen novum) to particular specimens or any contrary designation of type, or any different taxonomic usage of the new replacement name.
Type specimens deposited at the Zoological Museum Hamburg (ZMH) Superfamily Hepialoidea Stephens, 1829 Hepialoidea (the ghost moths and their relatives), with 629 described species worldwide, is the earliest diverging lepidopteran clade to exhibit any significant degree of species diversity today (Nielsen et al. 2000). The family Hepialidae is the most diverse group among other five families within the superfamily and comprises 606 species in 62 genera. In contrast to the remainder of the pre-Ditrysian Lepidoptera, many of the species are large and spectacular moths (Nielsen et al. 2000).   (Yakovlev 2011). The family Metarbelidae with more than 240 described species in 32 genera have their highest diversity in the Afrotropics (Lehmann 2019;Yakovlev and Zolotuhin 2020). However, the family is among the least known of Lepidoptera due their extreme rarity and rather cryptic appearance (Lehmann 2019 (Daniel 1932), he labelled all specimens (1♂ and 5♀) of the type series as cotypes; therefore all specimens are syntypes. Holotype (by original designation) in Museum Witt München, MWM (Yakovlev 2011). ZMH used to hold two syntypes of D. hethitica (Weidner 1974); however, the female specimen could not be located any more. Remarks. It was assumed that the syntypes were destroyed during the Allied bombing of Hamburg during World War II. However, as Warnecke reported to Daniel, two syntypes at the ZMH were lost in 1943, but one of them had been stored in Saxony and thus survived the bombings (Weidner 1974). Other type material (syntypes) is in the British Museum of Natural History (Yakovlev 2011;Yakovlev and Witt 2017).

Similipepsis osuni
The definition of the superfamily Zygaenoidea is based on only few characters. Although these characters are possibly autapomorphies this is partly not confirmed and is still a matter of discussion. The discussed characters are: head retractibility in larva, heteromorphosis, forward migration and reduction of head setae, position of microsetae and pores, labral sensilla (Vegliante and Zilli 2004). However, most of these characters have still not been sufficiently studied throughout all families. An overview regarding this discussion can be found in Efetov and Tarmann (2017). The Zygaenoidea contain the 13 families: Epipyropidae, Cyclotornidae, Himantopteridae, Anomoeotidae, Megalopygidae, Somabrachidae, Aididae, Limacodidae, Dalceridae, Lacturidae, Heterogynidae, Phaudidae, and Zygaenidae (Epstein et al. 1999;Efetov et al. 2014).
The family Zygaenidae has a world-wide distribution and is divided into the five subfamilies Inouelinae, Procridinae, Chalcosiinae, Callizygaeninae and Zygaeninae (Efetov and Tarmann 2017); it includes more than 1,000 species. 25 relevant characters that are possibly part of a 'hypothetical groundplan' of the family are discussed in detail in Efetov and Tarmann (2017). The most relevant possible autapomorphies of Zygaenidae are: 1. Cyanogenesis and resistance against cyanides. 2. Head with ocelli and chaetosemata present (as in some other more primitive ditrysian groups, e.g. Yponomeutoidea and Tortricoidea). This character combination is absent in all other Zygaenoidea. 3. Development of Petersen's gland (a pair of glands close to the ooporus in Zygaeninae and Procridinae of not exactly known function). This character is considered to be secondarily reduced in Inouelinae, Chalcosiinae and Callizygaeninae. This gland seems to have an importance for the protection of the eggs against predators.  (Hofmann and Tremewan 1996: 112).
Current status. Infrasubspecific and hence unavailable name.
Current status. Infrasubspecific and hence unavailable name.
Remarks. Reiss (1941) proposed this name as an aberration of Z. c. pinskica Reiss, 1941. Therefore, as stated by Articles 45.6.1 and 45.6.2 (ICZN 1999) it is deemed to be infrasubspecific name -if author used "aberration", "ab.", "morph", or the author expressly gave it infrasubspecific rank -which is hence unavailable.
Current status. Infrasubspecific and hence unavailable name.

Zygaena (Agrumenia) hilaris lucifera ab. gelpkei
Original locality. No type-locality mentioned in original description. This type comes from Sierra Alta de Albarracin.
Current status. Infrasubspecific and hence unavailable name.
Remarks. Reiss (1936) proposed this name as an aberration of Z. hilaris lucifera Reiss, 1936. Therefore, as stated by Articles 45.6.1 and 45.6.2 (ICZN 1999) it is deemed to be infrasubspecific name -if author used "aberration", "ab.", "morph", or the author expressly gave it infrasubspecific rank -which is hence unavailable.
Original locality. No type locality given for this taxon. This type comes from Sierra Alta de Albarracin. Current status. Infrasubspecific and hence unavailable name.
Original material examined. "Cotypes" 4♂♂ (ZMH 61490-ZMH 61493) (Fig. 50). "CALABRIA / bassingliatella oder Cassugliatella / 6.VIII.21 / ? // v. silaecola / Vrty // "Cotypus" / v. / silaecola / Vrty // Coll.  Schiffermüller, 1775). The taxon silaecola Verity, 1930 was proposed as an objective replacement name for Anthrocera meliloti silana Turati, 1923, which is a junior secondary homonym to Zygaena lonicerae silana Burgeff, 1914. However, the taxon silaecola Verity, 1930 is a junior synonym to sicula Calberla, 1895. The current combination of this taxon is therefore Z. viciae sicula Calberla, 1895 (Hofmann and Tremewan 1996). Moreover, as the reviewer Alberto Zilli correctly states, if the type status of specimen(s) in case an original name has objectively been replaced which is the case here as the name silaecola Verity, 1930 is stated to be a replacement for silana Turati, 1923 due to the latter entering homonymy with another name. In this case, ICZN article 72.7 clearly state that the type specimens remain those of the original name regardless of any statement to the contrary (e.g. other specimens named as types). Accordingly, the listed 'types' are such only if they are part of the original Turati's series, if they are part of another series worked upon by Verity they are not true types of silaecola. Consequently, the ‚'types' listed above are incorrectly labelled as types.
Current status. Valid subspecies.

Zygaena sarpedon leuzensis ab. quinquepuncta
Original locality. Not mentioned. The three syntypes are from the "Sierra Alta de Albarracin". Current status. Infrasubspecific and hence unavailable name.
Remarks. Reiss (1958) proposed this name as an aberration of Z. s. leuzensis Dujardin, 1956. Therefore, as stated by Article 45.6.1 (ICZN 1999) it is deemed to be infrasubspecific name -if its author expressly gave it infrasubspecific rank -which is hence unavailable.

Discussion
More than 2,000 type specimens of butterflies and moths (Lepidoptera) are deposited in the Zoological Museum of Hamburg (ZMH). In this catalogue, we have listed 259 "type" specimens: 13 valid species names, 34 subspecies (16 valid and 18 invalid names), and 36 invalid infrasubspecific rank under the ICZN code. Of these 259 specimens, 81 are primary types, 92 are secondary types, and 86 are considered invalid infrasubspecific rank (Suppl. material 1: Table S1).
Comparison of the first part of our updated catalogue to the type catalogue of Weidner (1974) reveals 32 new additions to the families Sesiidae, Brachodidae, Metarbelidae, and Zygaenidae (Suppl. material 1: Table S1). New additions to the type collection since the Weidner catalogue (1974) do not necessarily reflect "new type additions" -except for the five taxa donated by Kallies and Bartsch (Sesiidae and Brachodidae), and Lehmann (Metarbelidae) to the ZMH collection-they were found among the exisiting type material and were either mididentified or misplaced under the incorrect type labels. Details of the changes and updates including the new types, changes to the type status, and changes to the specimen counts since the last catalogue of Weidner (1974) were summarized in Suppl. material 1: Table S1.
The collection of Dr. Hanan Bytinski-Salz was the largest collection containing the most "type specimens" mainly from Hugo Reiss (89 specimens), Burchard Alberti (59 specimens) and Hanan Bytinski-Salz (35 specimens). None of the specimens from the latter author represents a valid name. The oldest type specimen was from the collection of Bytinski-Salz described by Korb in 1903(Zygaena rosinae Korb, 1903.