Illustrated catalogue of type specimens of Megalopodidae (Coleoptera) deposited at the Museu de Zoologia da Universidade de São Paulo

The Museu de Zoologia da Universidade de São Paulo (MZUSP) houses an important Megalopodidae collection representing 144 species, especially from Brazil and the Neotropical fauna. The type specimens of some species have never been accessed since their original descriptions, and are thus unknown to a major part of the Megalopodidae researchers. Presented here is an illustrated catalogue of the megalopodid type specimens of 32 species deposited at MZUSP, featuring photos of habitus and labels, and complete label data of all the specimens, which originally belonged to Jacintho Guérin’s personal collection. Conflicting and inconsistent data provided in the literature and in the specimens’ labels are addressed. Taxa originally proposed as species varieties are here reaffirmed as valid, with subspecific rank. Agathomerus varians Monrós, 1945 and Plesioagathomerus vittatus Monrós, 1945, originally described as junior synonyms, are considered unavailable names. The subspecific epithet ngriapex is emended into Agathomerus bifasciatus nigrapex Guérin, 1949. An overview of the MZUSP Megalopodidae collection is presented with a history of the arrival of Guérin’s specimens.


INTRODUCTION
Megalopodidae is a small family of Chrysomeloidea comprising over 580 species distributed worldwide, but with most of the diversity occurring in the Neotropical region (Lawrence & Ślipiński, 2014). Brazil is by far the most diverse country with about 150 species in 10 genera (Seeno & Wilcox, 1982;Rodríguez-Mirón, 2018). Despite the high diversity of species in Brazil, these numbers are expected to be underestimated due to the long-time absence of studies of the South American megalopodid fauna. After the contributions from Guérin (1943Guérin ( -1952, Monrós (1945Monrós ( , 1947 and Pic (e.g., 1948Pic (e.g., , 1955 in the decades of 1940 and 1950, no other taxonomic paper has been published about this family for the region. In addition, most of the Neotropical species are known only from their type-localities or from unclear or extensive localities (like Bolivia, Amazonia and South America) (e.g., Pic, 1955). Further studies in species distribution might show broader occurrence records and provide more accurate data on species richness and diversity.
The Megalopodidae collection from the Museu de Zoologia da Universidade de São Paulo (MZUSP, São Paulo, Brazil) stands out as one the most representative of the Neotropical region, especially from Brazil and neighbouring countries. It holds about 1,200 specimens in 144 species and seven genera, and other ca. 800 specimens still pending identification to species level. Besides, the MZUSP collection includes many type specimens and the totality or parts of important private collections, comprising specimens studied by several specialists in Chrysomeloidea (Campaner, 2014). For these reasons, the study of the MZUSP collection proves opportune for anyone assessing the Neotropical Megalopodidae fauna.
During a re-curation of the megalopodid collection, we came across specimens labelled as types of 30 taxa, some of which whose names do not figure in the recent world or local species lists (Rodríguez-Mirón, 2018;Botero, 2020, whereas others are cited only in the list of taxa from Brazil (Botero, 2020). These absences and conflicting data prompted us to investigate the validity of such names.
Some of these taxa were published in obscure journals with limited global reach, which are often unknown or overlooked by the coleopterist community. We therefore present an illustrated ISSN On-Line: 1807-0205 ISSN Printed: 0031-1049 ISNI: 0000-0004-0384-1825 and commented catalogue of the type specimens of Megalopodidae housed in the MZUSP in order to re-introduce these barely known species to science. The inconsistencies in the data provided in the literature and in the specimens' labels are commented and elucidated. Moreover, we address nomenclatural issues concerning the validity of taxa described as varieties and the availability of others originally described as synonyms. Finally, a brief account on the history of Monrós' and Guérin's collections and their arrival at the MZUSP.

MATERIAL & METHODS
All the specimens listed and illustrated are deposited in the Coleoptera collection of the Museu de Zoologia da Universidade de São Paulo (MZUSP). Information about complementary type series deposited in other collections is given when available.
The species are listed in alphabetical order in their original combination. A chronological list of citations is given for each entry and for their subsequent combinations and synonyms. The category of types (if holotype, paratype or syntype) is informed followed by a verbatim copy of labels' data. Data on different label rows are separated by a single slash ("/"), and data on different labels are separated by double slashes ("//"). When appropriate, conflicting data, corrections, justifications and further information on type series are commented.
Photographs of type specimens and their labels are provided for all the species, preferably the holotypes. Photographs were taken on a Zeiss AxioCam MRc 5 video camera attached to a Carl Zeiss Discovery stereomicroscope and edited on Adobe Photoshop CS6.

The arrival of the type specimens to the MZUSP
The major bulk of the MZUSP's Megalopodidae collection was formed in 1987 with the acquisition of the massive personal Coleoptera collection from Ricardo von Diringshofen ("Dirings") . Ferreira et al. (2016) presented biographical accounts and reported the processes of acquisition and incorporation of Dirings' entomological collection to the MZUSP. His collection was one of the most relevant of Brazil and the Neotropical Region especially in terms of the number of specimens, but also for comprising numerous other smaller collections purchased entirely or in part from Coleoptera specialists, such as J. Guérin, G.H. Nick, J. Bechyně, B. Pohl and F. Plaumann (Costa et al., 2000;Campaner, 2014;Campaner et al., 2008;Ferreira et al., 2016).
In the occasion of the arrival of Dirings collection to the MZUSP, Martins et al. (1991) presented a brief overview of the whole collection, then estimated in around 2.5 million specimens, and detailed the newcomer chrysomeloid collection in each subfamily, including the number of types, total number of specimens, and those identified by Chrysomeloidea specialists, such as J. and S. Bechyně, W. Bokermann, J. Guérin and E. Uhmann.
The incorporation of Dirings' collection to the MZUSP brought not only a great number of specimens of Megalopodidae originally assembled by him and his collaborators, but also parts of the excellent collection previously acquired from Jacintho Guérin. Dirings' megalopodid holdings comprised 776 specimens, 617 of which determined by specialists, notably F. Monrós and J. Guérin (Martins et al., 1991).
Jacintho Guérin (1898Guérin ( -1960 was an Italian engineer who lived most of his life in Brazil, where he worked for an electricity distribution company (Nomura, 1995). He started to form his entomological collection in 1922 with butterflies, but soon after gave priority to beetles.
His collection comprising around 30,000 specimens is now deposited in the Instituto Biológico de São Paulo (IBSP, São Paulo, Brazil), except for those from the families Erotylidae, Megalopodidae and Chrysomelidae: Cryptocephalinae, which had previously been acquired by Dirings (Nomura, 1995;Campaner et al., 2008). Every specimen in his collection received an accession number that would be mentioned in his papers. Such numbers enable a reliable recognition of the specimens when these are mislabelled in the collection. We could not retrieve his record book, which might possibly be held by the IBSP. Originally, within each drawer, the species were organised according to the colour patterns and similarities, thus enabling their easy recognition and identification. Once arrived at the MZUSP, his collection was disassembled and the species scattered along with the MZUSP's megalopodid collection and displayed in alphabetic order.
Guérin's collection included types and other specimens originally from Francisco Monrós, an author with whom he was in constant contact and eventually exchanged material. Francisco Monrós (1922Monrós ( -1958) was a prolific Spanish taxonomist working in Argentina specialized in the smaller Chrysomeloidea families and subfamilies. During his short career, he assembled a substantial collection that included specimens exchanged with many eminent colleagues of his time. Upon his death, the collection was acquired by the United States National Museum (Washington, D.C., USA). Biographies and detailed information about his work and collection can be found in Blake (1958Blake ( , 1961 and Staines (1995).
We call the attention to the fact that some type specimens from Guérin's collection might not be labelled accordingly. Specimens of Agathomerus nicki Guérin, 1948 and Nickimerus setosus Guérin, 1948, labelled as allotypes, are actually the holotypes. One specimen labelled as cotype (= syntype) of Plesioagathomerus canus Monrós, 1945 actually does not belong to the type series, since it has been collected after the species description. Other mislabelled specimens are herein corrected and commented on. Furthermore, the species that were recognized by Guérin as junior synonyms (either published or not) received a new identification label, had their red type labels removed, and were organized under the name of the senior synonym. For example, the holotype of Agathomerus nobilis cyaneonotatus was displayed amongst specimens of their senior synonym A. basalis Pic, 1916, without a red type label. Moreover, the types of Mastostethus semirufus Guérin, 1943 andMegalopus vittatus Guérin, 1943 were found amongst specimens of Mastostethus angustalisi Pic, 1916 andMegalopus unifasciatus Pic, 1916, respectively, albeit these species have not been formally synonymized. A similar organisation fashion is expected to be found in other parts of Guérin's collection, in which the putative "lost" types might be retrieved after a scrutiny throughout the collection, and they can be recognized by the matching data and accession numbers given in the original description and the specimens' labels. Guérin, 1946 ( Fig. 1A-C) Agathomerus almeidai Guérin, 1946: 205, fig. 1

Comments:
The original description clearly mentions the black spots in the apex of elytra as a diagnostic character for the subspecies. Thus, the spelling "ngriapex" is here considered a lapsus, and "nigrapex" is accepted as the intended correct name. The taxon name is therefore corrected to Agathomerus bifasciatus nigrapex Guérin, 1949, nomem emendatum (ICZN, 1999.