TY - JOUR
AU1 - Ferreira, Vinicius S
AU2 - Ivie, Michael A
AB - Abstract The Leptolycini are a group of Lycidae endemic to the West Indies. Leptolycini adult females have been hypothesized to be extreme paedomorphic (i.e., larviform), however, females and larvae of the group are currently unknown. Here we provide the first association of adult male and immature life stages from the Puerto Rico using DNA barcoding, also collections-based associations and descriptions of immature Leptolycini and the first description of a paedomorphic female from the Virgin Islands. To carry out these life-stage associations we prepared an in-depth review of the Leptolycini fauna of the Puerto Rican bank (Puerto Rico and Virgin Islands). Several new taxa and taxonomic arrangements are proposed: Cessator crypticusnew species, Cessator tortolensisnew species, Cessator obrienorumnew species; Dracolycus chupacabranew genus and species, Dracolycus marshallinew species; Leptolycus falsoheterocornisnew species, and Leptolycus viensisnew species. Nanolycus gnomus Kazantsev is moved to Cessator gnomus (Kazantsev) new combination, rendering Nanolycus Kazantsev a new junior synonym of Cessator Kazantsev. The subgenus Baholycus Bocak is a new junior synonym of Leptolycus Leng and Mutchler. Leptolycus heterocornis var. flavicollis Leng and Mutchler is elevated to Leptolycus flavicollis Leng and Mutchler new status; Leptolycus (Leptolycus) albicauda Kazantsev is a new junior synonym of Leptolycus flavicollis Leng and Mutchler. An updated key to the adults and immature forms of Leptolycini from the Puerto Rican bank and a discussion on the importance of scientific collections in biodiversity studies is also provided. The Leptolycini are a group of Lycidae endemic to the West Indies for which only the males are known and described. The morphology of the Leptolycini adult males conforms with other groups suspected or proven to be affected by the paedomorphosis syndrome, and females of this tribe are hypothesized to be larviform. Throughout the years, many different classifications proposed various taxonomic rank and tribal and generic arrangements for the members of the Leptolycini (see competing classification systems in Bocak and Bocakova 1990, Kazantsev 2005, Bocak et al. 2008, Kazantsev 2013, 2017; Bocakova 2014). Despite the controversial classification status of the Leptolycini, the most recent molecular-based phylogenetic hypotheses for Lycidae supported the tribe as an independent lineage endemic to the West Indies (Masek et al. 2018, Kusy et al. 2019). Leptolycini, as currently understood, is comprised by six described genera: CessatorKazantsev 2009, DominopteronKazantsev 2013, ElectropteronKazantsev 2012, LeptolycusLeng and Mutchler 1922, NanolycusKazantsev 2013, and TainopteronKazantsev 2009, and a total of 20 described species (Leng and Mutchler 1922, Zayas 1988, Bocak 2001, Kazantsev 2013; 2017, Ferreira and Ivie 2017). The tribe contains many other undescribed genera and species, and a revision of the group is currently in preparation by Ferreira and Ivie (in prep). Recent molecular phylogenies (Masek et al. 2018, Kusy et al. 2019) indicated that many of the taxa previously placed in the Leptolycini are now understood as belonging to the mainly mainland Calopterini (Lycinae) (such as Lycinella Gorham, 1884, Cephalolycus Pic, 1926, and Aporrhipis Pascoe, 1887, and all other Neotropical lycids suspected to be paedomorphic) (e.g., Bocakova 2003, 2005; Kazantsev 2005, 2013, 2017; Ferreira and Ivie 2016, 2017, 2018; Ferreira 2020; Ferreira and Silveira 2020) and dispersed in other lycid subfamilies (see additional references below). Paedomorphosis (PDM) is a type of heterochrony in which the development of certain traits is truncated relative to an organisms’ ancestral form (Reilly et al. 1997, McNamara 2012). In animals, PDM is a syndrome in which juvenile traits of an organism are retained into adulthood (Reilly et al. 1997, McNamara 2012). Beetles affected by the paedomorphic syndrome generally have reduced sclerotization and flight ability, miniaturized structures, and the predominance of a K-reproductive strategy (Bocakova et al. 2007, Bocak et al. 2008, McMahon and Hayward 2016). In some lineages, individuals possess a mosaic of combinations between fully developed adult individual and immature features. These can be expressed by an adult-like head and prothorax and a distended abdomen (Lampyridae, Elateridae: Omalisinae, Plastocerinae and Drilinae) (Cicero 1988, Bocek et al. 2018, Kusy et al. 2018a), or distended abdomen with reduced elytra and less sclerotized (soft) bodies (e.g., Dermestidae, Staphylinidae, Micromalthidae, Lampyridae, Elateridae: Cebrionini, Staphylinidae) (Mertins 1981, Cicero 1988, Kundrata and Bocak 2007, Perotti et al. 2016, Kusy et al. 2018a, Rattu 2020, Zilberman 2020). To the extreme, certain adult females strongly resemble their immature stages, and these forms are in the literature usually referred to as “larviform females” (e.g., Dermestidae, Lycidae, Phengodidae) (e.g., Mertins 1981, Masek and Bocak 2014, Roza et al. 2017, Kundrata et al. 2019). In Coleoptera, the PDM syndrome occurred independently multiple times in several unrelated lineages (McMahon and Hayward 2016, Vea and Minakuchi 2021), with a higher preponderance of cases within the superfamily Elateroidea (Crowson 1972, Cicero 1988, Bocakova et al. 2007, Kundrata et al. 2014), which includes some of the most popular and charismatic beetle groups, such as fireflies (Lampyridae) (e.g., Martin et al. 2017, 2019; Ferreira et al. 2019, 2020), click-beetles (Elateridae) (e.g., Rattu 2016, Kundrata et al. 2019, Zapata de la Vega et al. 2020), rail-road worms (Phengodidae) (e.g., Roza et al. 2017), and the so called “trilobite-larvae”, and other net-winged beetles (Lycidae) (e.g., Masek and Bocak 2014, Kazantsev 2017, Ferreira and Silveira 2020), to mention a few known or suspected cases of paedomorphic lineages in this superfamily. Within the Elateroidea, PDM has evolved multiple times, and recent molecular-based phylogenetic hypotheses indicate that PDM has happened at least five times independently in Lycidae, in the Lyropaeinae, Ateliinae, Lycinae, Metriorrhynchinae, and Dexorinae (Fig. 1A) (sensuKusy et al. 2019) (Bocak and Bocakova 2008, Masek et al. 2018, Kusy et al. 2019). Fig. 1. Open in new tabDownload slide A. A scheme showing the hypothesized evolutionary relationships of Lycidae (modified from Kusy et al. 2019). Red circles indicate groups in which paedomorphosis has developed at least once in the subfamilies. B. Distribution of Lycidae lineages across the different zoogeographical zones. Map of the different ecozones modified from Wikimedia commons. Originally made by carol and used under CC BY-SA 3.0. Fig. 1. Open in new tabDownload slide A. A scheme showing the hypothesized evolutionary relationships of Lycidae (modified from Kusy et al. 2019). Red circles indicate groups in which paedomorphosis has developed at least once in the subfamilies. B. Distribution of Lycidae lineages across the different zoogeographical zones. Map of the different ecozones modified from Wikimedia commons. Originally made by carol and used under CC BY-SA 3.0. In Lycidae, paedomorphic modifications in the adult male’s morphology are often associated with a general miniaturization process and an overall less sclerotized integument (i.e., softbodiness) (Miller 1991, Bocakova 2003, 2005; Bocak et al. 2008, Bocakova 2014, Masek and Bocak 2014, Masek et al. 2014, Bocak 2018, Ferreira and Ivie 2018, Ferreira 2020). The mouthparts are often reduced in size or completely or partially absent, and the number of antennomeres might be reduced in certain genera (Kazantsev 2009, 2013, 2017; Bocakova 2014, Masek and Bocak 2014, Masek et al. 2014, Bocak 2018, Ferreira and Ivie 2018, Ferreira 2020). The wings might present reduced venation and the elytra, which are noncoadapted with the abdomen and thorax and are frequently dehiscent, often present a reduced number or complete absence of transversal and longitudinal costae and might be strongly punctate (Miller 1991, Bocakova 2003, 2005; Bocak et al. 2008, Kazantsev 2009, 2013, 2017; Bocakova 2014, Masek and Bocak 2014, Masek et al. 2014, Bocak 2018, Ferreira and Ivie 2018, Ferreira 2020). The abdomen might be physogastric and the genitalia simplified when compared to other Lycidae (Miller 1991, Bocakova 2003, 2005; Bocak et al. 2008, Kazantsev 2009, 2013, 2017; Bocakova 2014, Masek and Bocak 2014, Masek et al. 2014, Bocak 2018, Ferreira and Ivie 2018, Ferreira 2020). Several lycid groups for which the males normally present different degrees of the above-mentioned characteristics females are known to be paedomorphic (see below), but in most cases females are completely unknown and only presumed to be paedomorphic. Groups like this include several Calopterini and Leptolycini (Lycinae) and the Dexorinae and Ateliini (Ateliinae)) (e.g., Kazantsev 2005, Bocakova 2014, Kazantsev 2017, Li et al. 2018a, Ferreira and Silveira 2020). In other groups, such as the Lyropaeinae, adult females are known to be extreme paedomorphic (Mjöberg 1925, Miller 1991, Wong 1996, Masek et al. 2014). These females are often referred to as “neotenous”, “pedogenic”, or “larviform” (Mjöberg 1925, Wong 1996, Miller 1991, Bocakova 2003, 2005; Bocak et al. 2008, Kazantsev 2009, 2013, 2017; Bocakova 2014, Masek and Bocak 2014, Masek et al. 2014, Bocak 2018, Ferreira and Ivie 2018, Ferreira 2020). Even though PDM is a widely spread phenomenon in Lycidae, very little is known in the literature about the morphology of extreme paedomorphic females and other male-female associations in other paedomorphic lineages in the family. In the Lyropaeinae, the extreme paedomorphic females of Platerodrilus spp. have been described by Mjöberg (1925), Wong (1996), and Bocak and Matsuda (2003), whilst DNA-based associations of males and females were made by Levkanikova and Bocak (2009) and Masek and Bocak (2014). Adult-larvae and male-female associations were also provided in Masek et al. (2014) for Lyropaeus sp. DNA-based associations of adults and larvae were also provided by Levkanikova and Bocak (2009) for Macrolibnetis depressus Pic, 1938. In the Lycinae, except for the descriptions and the tentative association of larval/extreme paedomorphic female stages of Pseudacroleptus lamelliferKazantsev and Zaitsev, 2008 and Ceratoprion ignavumKazantsev and Zaitsev, 2008 (Calopterini), nothing else has been published on this for any other New World Lycidae lineage affected by PDM (Kazantsev and Zaitsev 2008). To further contribute to the topic, in this study we document the first associations of extreme paedomorphic female individuals with their immature stages and respective males in beetles of the tribe Leptolycini. VSF’s recent collection trip to Puerto Rico and MAI’s extensive collection of specimens and biological observations of beetles from the West Indies allowed the authors to provide first-hand DNA barcoding associations based on partial cytochrome c oxidase subunit 1 (COI) of winged males and immature stages of specimens from Puerto Rico, together with collections-based associations of Leptolycini immatures and one extreme paedomorphic female from the Virgin Islands. However, due to the lack of taxonomic knowledge of the Lycidae fauna of the region, to carry out these life-stage associations we had to prepare an in-depth review of the Leptolycini fauna of the Puerto Rican bank (Puerto Rico and Virgin Islands), which resulted in many taxonomic and nomenclatural changes and the descriptions of several new taxa. We provide morphological diagnoses of the larvae and female that were found, diagnostic redescriptions of the already known adult males and full descriptions of the new ones. An updated key to the adults and immature forms of Leptolycini from the Puerto Rican bank and a discussion on the importance of scientific collections in biodiversity studies, especially regarding understudied groups, like the Leptolycini, is also provided. Material and Methods Study Areas The material used in this study is the collective efforts of dozens of collectors and sampling efforts from collection expeditions to Puerto Rico and to the Virgin Islands (Fig. 2A–C), with most of the specimens coming from the El Verde Field Station (EVFS) in Puerto Rico (Fig. 2B). Puerto Rico (PR) is the place with the highest known diversity of Leptolycini in a single locality, with seven species of three genera known from the males (Leng and Mutchler 1922, Kazantsev 2009, 2013) and several morphotypes of larvae/females occurring at the EVFS in the El Yunque National Forest in the Luquillo Mountains and/or the surrounding Caribbean National Forest in Puerto Rico (Fig. 2B), with another two species known from males elsewhere in PR. The EVFS is located 10 Km south of Rio Grande at 500 m above the sea level, with mean monthly temperatures ranging from 21°C in January to 25°C in September, and an annual precipitation average of 370 cm, with 35–40 cm per month in the wet season (May–December) (Brown et al. 1983, McDowell et al. 2012, Schowalter et al. 2014 and references therein). The area is cojointly administered by the Department of Environmental Sciences, University of Puerto Rico, Río Piedras, and the United States Department of Agriculture (USDA) Forest Service, and is the only location within the United States, other than Hawaii, with a tropical rain forest ecosystem (EVFS 2020). Fig. 2. Open in new tabDownload slide Study areas of the sampled material. A. Position of the Puerto Rican bank in the West Indies. B. Locality of the El Verde Field Station in the El Yunque National Forest in Puerto Rico. C. The Virgin Islands in detail; named islands are the ones in which Leptolycini specimens were collected. Fig. 2. Open in new tabDownload slide Study areas of the sampled material. A. Position of the Puerto Rican bank in the West Indies. B. Locality of the El Verde Field Station in the El Yunque National Forest in Puerto Rico. C. The Virgin Islands in detail; named islands are the ones in which Leptolycini specimens were collected. Specimens Sampling, Identification, Photographs, Maps, and Illustrations Sampling of DNA grade specimens was conducted in June of 2016 in Puerto Rico by Martin Fikáček and collaborators (Národní Museum, Czechia), in June and July of 2017 in the EVFS by VSF and Erich Spiessberger, and in July of 2018 by T. Sanchez & K. Noa Reyes and other students in a project coordinated by Donald Yee (University of Southern Mississippi). Additional non-DNA-grade specimens from the EVFS were collected by Eduardo Nazaro between May and November of 1996, whilst material from other places in the Virgin Islands have been collected by MAI and collaborators throughout the past 40 y during many collecting expeditions. Specimens were collected using a variety of methods: sweep-netting (adult males), by hand (males, females, and larvae), Malaise-traps and Flight Intercept Traps (FIT) (males, females, and larvae) (Fig. 3), soil washing, Berlese/Winkler extractions from leaf litter and moss-leaf-litter samples and pitfalls (females and larvae). Transcription of label data from specimens is as follows: the end of each line on a label is indicated by a “;” (semicolon); the individual labels are separated by a “/” (slash). Examined specimens are or will be deposited in the following collections (curators names presented after the institution locality): Fig. 3. Open in new tabDownload slide Puerto Rico collection site and some of the used collecting methods. A. Tropical rainforest in the El Verde Field Station. B. Flight interception trap (FIT). C. Malaise trap. D. Leptolycini specimens (above Leptolycus puellus and to the right Cessator luquillonis) first collected by net-sweeping and then with an aspirator. Fig. 3. Open in new tabDownload slide Puerto Rico collection site and some of the used collecting methods. A. Tropical rainforest in the El Verde Field Station. B. Flight interception trap (FIT). C. Malaise trap. D. Leptolycini specimens (above Leptolycus puellus and to the right Cessator luquillonis) first collected by net-sweeping and then with an aspirator. AMNH—American Museum of Natural History, New York, New York, U.S. (Lee Herman Jr.). CNC—Canadian National Collection of Insects, Ottawa, Ontario, Canada (Hume Douglas). DEBU—University of Guelph, Guelph, Ontario, Canada (Steve Marshall). EPRL—University of Puerto Rico, Mayaguez, Puerto Rico (Rafael Ingles). MLGC—Matthew L. Gimmel Private Collection, Santa Barbara, California, U.S. MCZ—Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, U.S. (Crystal Maier). NCSU—North Carolina State University Insect Collection, Raleigh, North Carolina, U.S. (Bob Blinn). NMNH—National Museum of Natural History, Washington D.C., U.S. (currently at the Montana Entomology Collection, Montana State University, Michael A. Ivie). NMPC—National Museum, Prague, Czechia f(Lukáš Sekerka). WIBF—West Indian Beetle Fauna Project Collection, Montana State University, Bozeman, Montana, U.S. (Michael A. Ivie). Specimens were sorted to morphospecies under a Leica Wild M3C stereoscopic microscope with magnification up to 40× and identified using available literature (Leng and Mutchler 1922, Kazantsev 2009, 2013, 2017) and by comparison of material with type specimens. Individuals were initially grouped based on morphological similarity and a temporary numeric code was assigned to each morphotype for which representative specimens were imaged. Adult morphological terminology followed Bocak and Bocakova (1990), Kazantsev (2005), and Lawrence et al. (2011) and immatures and larviform female terminology follows Bocak and Matsuda (2003), Levkanikova and Bocak (2009) and Ferreira and Costa (2015). Lycidae larvae and extreme paedomorphic females (lacking compound eyes, bifid tarsal claws, a multi-piece antennal flagellum and elytra) can be usually distinguished from other Elateroidea by the nonopposable mandibles that are each divided longitudinally, and the circular, weakly developed pygopodium in Lycidae (vs. a several finger-like, strongly developed structure in the Lampyridae). Photos of specimens for illustrations were taken using a Canon T3i or 6D DSLR with the MP-E 65 mm lens and using a Stackshot—automated macro rail for focus stacking and a JVC (DC Ky-F75U) digital camera mounted on a Leica MS5 stereoscope. Specimens preserved in ethanol were first placed in a container with dH2O for at least five minutes and posteriorly positioned for images in a water-based lubricating gel that was fulfilled with dH2O. Images were then stacked using Zerene Stacker version 1.04 and enhancements to digital images were made in Adobe Photoshop CC 2020 and Adobe Photoshop CC for iPad, while plate preparations were made in Adobe Illustrator CC 2020. The distribution maps were prepared using the software Quantum Gis 3.16. The distribution points were based on label information of examined specimens and georeferenced using Google Earth, exported to the format.kml and imported to Quantum Gis. The used raster files were downloaded from the following repositories: https://data.humdata.org/, https://data.fs.usda.gov/geodata/edw/datasets.php and https://luq.lter.network/data/luqmetadata9/5872. DNA Extraction and Amplification Settings We extracted whole genomic DNA using the Qiagen DNeasy Blood and Tissue Kit (Qiagen, Germantown, MA) from whole specimens (Genbank accession codes MW560932–MW560954, see full list in Supp. Material S1 [online only]). Extractions were permanently deposited in the MTEC DNA collection, at –80°C; after curation, adult specimens were either remounted and returned to the dry collection or were permanently deposited in vials in EtOH 70% in the immature beetles’ collection of WIBF in MTEC. PCR amplifications were performed with 25 µl reaction volume, using 12.5 µl GoTaq Green Master Mix (Promega, Madison, WI), 6.5 of UltraPure DNase/RNase-Free Distilled Water, 2 µl of each primer, and 2 µl of template to amplify the “barcode” region of cytochrome oxidase subunit I gene (COI). The polymerase chain reaction (PCR) and primers (LCO and HCO) conditions followed Folmer et al. (1994) and Raupach et al. (2016). PCR amplicons were sequenced in both directions by MCLAB (San Francisco, CA). Data Analyses, Genetic Distances Calculations, and Biogeography-Based Association Newly produced sequences were assembled in Geneious 11.1.2 (Biomatters Ltd., Auckland, New Zealand) and individually examined for any discrepancies and irregularities. Alignments were performed using MAFFT 7 under the G-INS-i algorithm for highly conserved sequences implemented in the MAFFT server (https://mafft.cbrc.jp/alignment/server/) (Katoh et al. 2002). Pairwise genetic distances calculations were implemented in MEGA 10.0.8 (Kumar et al. 2018) and in Geneious, using the uncorrected p-distances (PD) (Fitch and Margoliash 1967), Kimura’s 2-parameter (K2P) test (Kimura 1980), the Jukes-Cantor (JC) (Jukes and Cantor 1969), and the percentage identity (PC) criterion methods. We used the threshold percentages for species delimitations recommended in Ratnasingham and Hebert (2013) and followed by Raupach et al. (2016) for intraspecific (≤ 2.2%) and interspecific (≥ 2.3%) distances. For graphical representations of the genetic distances, we performed a maximum-likelihood analysis with 5,000 bootstraps using W-IQTREE (Trifinopoulos et al. 2016). A sequence of an individual identified as Calopteron reticulatum (Fabricius 1775), a distantly related Lycidae taxon, was downloaded from Genbank (voucher JN288183) and used to root the tree. Further life stage associations and male-female associations were made based on information from collections-based specimens and their known distribution patterns. In our study, some of the taxa in the Virgin Islands are only known to occur in specific localities, from which no other Leptolycini or even other Lycidae species are known, and scientific collection-based analyses of specimens sampled for these localities allowed us to infer associations for these organisms. Nomenclature This paper and the nomenclatural act(s) it contains have been registered in Zoobank (www.zoobank.org), the official register of the International Commission on Zoological Nomenclature. The LSID (Life Science Identifier) number of the publication is: urn:lsid:zoobank.org:pub:E8A5D76B-93A2-482E-A368-2F1F2FFF1BC Results Puerto Rican Samples Identification, COI Barcode Production, and DNA Based Associations From our sampling efforts of adult males from Puerto Rico, we identified 12 species belonging to four genera. Of these genera, three were already known and described: Cessator, Leptolycus, and Tainopteron, whilst one (Dracolycusnew genus) is herein described for the first time. Of the 12 species encountered, we were able to find and study specimens of all five previously known species for the Puerto Rican bank: Cessator luquinollisKazantsev, 2009, Leptolycus albicaudaKazantsev, 2009, Leptolycus puellusKazantsev, 2009, Leptolycus heterocornisLeng and Mutchler, 1922, and Tainopteron milleriKazantsev, 2009; the remaining seven species are described here for the first time (three in Cessator, two in Dracolycus and two in Leptolycus). From our sampling of immature/females, we initially identified 12 morphotypes that were later narrowed down to nine (nine larval type and one extreme paedomorphic female were identified). Of this material, we were able to amplify and sequence COI sequences of five male adult species (n = 10) and five larvae (n = 13) (see Table 1). Table 1. Adult males and immature forms for which COI sequences were amplified Taxon/Morphospecies of Leptolycini . DNA vouchers . Males Cessator tortolensis MTEC 023637 Cessator luquinollis WIBF 051525 Cessator crypticus MTEC 023638 Leptolycus puellus WIBF 041665, MTEC 023635, WIBF 051534 WIBF 051533, Leptolycus heterocornis WIBF 041667, MTEC 023636, WIBF 051532 Immature forms Leptolycus heterocornis WIBF 041693, WIBF 051506, WIBF 051507, WIBF 051537 Cessator luquillonis WIBF 041694, WIBF 051497, WIBF 051498 Cessator crypticus WIBF 041695, WIBF 041696, MTEC 023648 Dracolycus sp. 1 MTEC 023640, WIBF 051523 Dracolycus sp. 2 WIBF 041511 Taxon/Morphospecies of Leptolycini . DNA vouchers . Males Cessator tortolensis MTEC 023637 Cessator luquinollis WIBF 051525 Cessator crypticus MTEC 023638 Leptolycus puellus WIBF 041665, MTEC 023635, WIBF 051534 WIBF 051533, Leptolycus heterocornis WIBF 041667, MTEC 023636, WIBF 051532 Immature forms Leptolycus heterocornis WIBF 041693, WIBF 051506, WIBF 051507, WIBF 051537 Cessator luquillonis WIBF 041694, WIBF 051497, WIBF 051498 Cessator crypticus WIBF 041695, WIBF 041696, MTEC 023648 Dracolycus sp. 1 MTEC 023640, WIBF 051523 Dracolycus sp. 2 WIBF 041511 Open in new tab Table 1. Adult males and immature forms for which COI sequences were amplified Taxon/Morphospecies of Leptolycini . DNA vouchers . Males Cessator tortolensis MTEC 023637 Cessator luquinollis WIBF 051525 Cessator crypticus MTEC 023638 Leptolycus puellus WIBF 041665, MTEC 023635, WIBF 051534 WIBF 051533, Leptolycus heterocornis WIBF 041667, MTEC 023636, WIBF 051532 Immature forms Leptolycus heterocornis WIBF 041693, WIBF 051506, WIBF 051507, WIBF 051537 Cessator luquillonis WIBF 041694, WIBF 051497, WIBF 051498 Cessator crypticus WIBF 041695, WIBF 041696, MTEC 023648 Dracolycus sp. 1 MTEC 023640, WIBF 051523 Dracolycus sp. 2 WIBF 041511 Taxon/Morphospecies of Leptolycini . DNA vouchers . Males Cessator tortolensis MTEC 023637 Cessator luquinollis WIBF 051525 Cessator crypticus MTEC 023638 Leptolycus puellus WIBF 041665, MTEC 023635, WIBF 051534 WIBF 051533, Leptolycus heterocornis WIBF 041667, MTEC 023636, WIBF 051532 Immature forms Leptolycus heterocornis WIBF 041693, WIBF 051506, WIBF 051507, WIBF 051537 Cessator luquillonis WIBF 041694, WIBF 051497, WIBF 051498 Cessator crypticus WIBF 041695, WIBF 041696, MTEC 023648 Dracolycus sp. 1 MTEC 023640, WIBF 051523 Dracolycus sp. 2 WIBF 041511 Open in new tab The length of the generated DNA barcode fragments ranged from 548 to 667 bp and our sequence data revealed a high AT-content, with mean sequence compositions of A = 35.7%, C = 18.3%, G = 12.6%, T = 33.4% (aligned sequences available in Supp. Material S2 [online only]). The different methods we used for association of specimens (PD, K2P, JC, PC) rendered nearly identical results (Supp. Materials S3–S6 [online only] for PD, K2P, JC, PC, respectively). With our DNA analyses, we were able to provide matches for the following species: C. luquillonis, C. crypticus, and L. heterocornis (Fig. 4) (See Supp. Materials S3–S6 [online only] for PD, K2P, JC, PC). Using the PC method, intraspecific distances of 0.53% or less were found for the L. heterocornis adult and its larvae. For C. luquillonis adult and its respective larva match, we found intraspecific distances of 1.04%, whilst for C. crypticus adult male and its larvae, we found intraspecific distances of 0.17% or less. No matches were found for L. puellus, C. tortolensis, and what we are considering to be the larvae of Dracolycus (see discussion under descriptions and diagnosis of that genus), for which their respective life-stage counterparts remain unknown. Interspecific differences ranged from ~25% to ~13%, with the smallest differences found between congeneric taxa (L. puellus and L. heterocornis) (see full numbers in Supp. Materials S3–S6 [online only] for PD, K2P, JC, PC). Fig. 4. Open in new tabDownload slide Maximum-likelihood topology of the generated COI sequences and a mainland Calopteron. Branches in color represent matches between adult males and larvae. Numbers represent SH-aLRT (left) and Ultrafast Bootstrap (UFboot) (right). Fig. 4. Open in new tabDownload slide Maximum-likelihood topology of the generated COI sequences and a mainland Calopteron. Branches in color represent matches between adult males and larvae. Numbers represent SH-aLRT (left) and Ultrafast Bootstrap (UFboot) (right). The graphical display of the relationships of the sequenced taxa are presented in the form of a tree topology (Fig. 4 and Supp. Material S7 [online only] for the support level of all included taxa). Our ML tree recovered a cluster with males and larvae of L. heterocornis as sister to L. puellus (Fig. 4). The Leptolycus cluster was recovered in a polytomy that contained a cluster for C. luquillonis adult and larvae, C. crypticus and larvae, Dracolycus sp. 1 + Dracolycus sp. 2, and C. tortolensis by itself (Fig. 4). Biogeography Life-Stages Association in the Virgin Islands Due to the well-studied beetle fauna of the Virgin Islands and the cross-matching of results with our DNA-based life stages association from Puerto Rico, we were able to confidently provide association of life stages of the Leptolycini specimens found in the Virgin Islands. From our sampling efforts of adult Leptolycini males from the Virgin Islands, we identified two species belonging to two genera. We found a new species of Leptolycus distributed in St. John, Guana and Tortola (Fig. 2C) and two morphotypes of immature forms/larviform females associated with these samples from St. John (Fig. 2C). One new species of Cessator was found in Tortola together with its larval form (Fig. 2C). The larvae and extreme paedomorphic females of Leptolycus collected in St. John are the results of several expeditions (see details in the Taxonomic Revision in the next section), while the larvae (and all known adults) of immature Cessator from Tortola are only known from a single collecting event. These new species and their respective semaphoronts are described below and diagnostic characters to separate larvae from females are presented as well. Taxonomic Revision of the Puerto Rican Bank Leptolycini Fauna To carry out our life-stage associations on the Puerto Rican bank Leptolycini and to be able to correctly identify the taxa from the region, we needed to properly know the region’s faunal diversity. Using published literature and with access to materials from several collections, including type specimen vouchers or photographs (see material and methods section), we were able to provide an in-depth review of the Leptolycini fauna of the Puerto Rican bank. Short diagnostic descriptions and illustrations of the previously known taxa are provided, as well as full descriptions of the new ones. Diagnosis of the immature forms and of the female are provided for the first time. Identification keys for the adult male genera, the species in each genus and to the known immature and extreme paedomorphic females of Leptolycini from the Puerto Rican bank are also provided. These are provided below, presented in alphabetical order for the genera and in publication chronological order for the species within each genus. CessatorKazantsev, 2009 Figs. 4, 5A–D, 6A–D, 7A–D, 8A–C, 9A–B, 10A–C, 11, 12A Fig. 5. Open in new tabDownload slide Male adult habiti of the Puerto Rican bank Leptolycini—Cessator, Dracolycus and Tainopteron. A. Cessator luquillonisKazantsev 2009. B. Cessator tortolensisnew species. C. Cessator obrienorumnew species. D. Cessator crypticusnew species. E. Dracolycus chupacabranew genus and new species. F. Dracolycus marshallinew genus and new species. G. Tainopteron milleriKazantsev 2009. Fig. 5. Open in new tabDownload slide Male adult habiti of the Puerto Rican bank Leptolycini—Cessator, Dracolycus and Tainopteron. A. Cessator luquillonisKazantsev 2009. B. Cessator tortolensisnew species. C. Cessator obrienorumnew species. D. Cessator crypticusnew species. E. Dracolycus chupacabranew genus and new species. F. Dracolycus marshallinew genus and new species. G. Tainopteron milleriKazantsev 2009. Fig. 6. Open in new tabDownload slide Head ventral detail of the adult male Puerto Rican bank Leptolycini—Cessator, Dracolycus and Tainopteron. A. C. luquillonis. B. C. tortolensis. C. C. obrienorum. D. C. crypticus. E. D. chupacabra. F. D. marshalli. G. T. milleri. Fig. 6. Open in new tabDownload slide Head ventral detail of the adult male Puerto Rican bank Leptolycini—Cessator, Dracolycus and Tainopteron. A. C. luquillonis. B. C. tortolensis. C. C. obrienorum. D. C. crypticus. E. D. chupacabra. F. D. marshalli. G. T. milleri. Fig. 7. Open in new tabDownload slide Male genitalia of Leptolycini species, dorsal view. Cessator spp. A–D. A. C. luquillonis. B. C. tortolensis. C. C. obrienorum. D. C. crypticus. Dracolycus spp. E–F. E. D. chupacabra. F. D. marshalli. G. T. milleri. Leptolycus spp. H–N. H. L. flavicollis. I. L. viensis. J. L. falsoheterocornis. K. L. puellus. L. heterocornis color morphs L–N. L. Color morph 1. M. Color morph 2. N. Color morph 4. Fig. 7. Open in new tabDownload slide Male genitalia of Leptolycini species, dorsal view. Cessator spp. A–D. A. C. luquillonis. B. C. tortolensis. C. C. obrienorum. D. C. crypticus. Dracolycus spp. E–F. E. D. chupacabra. F. D. marshalli. G. T. milleri. Leptolycus spp. H–N. H. L. flavicollis. I. L. viensis. J. L. falsoheterocornis. K. L. puellus. L. heterocornis color morphs L–N. L. Color morph 1. M. Color morph 2. N. Color morph 4. Fig. 8. Open in new tabDownload slide Dorsal view of the immature forms of the Puerto Rican bank Leptolycini. Cessator and Dracolycus larvae spp. A–C. Cessator spp. A. C. crypticus. B. C. luquillonis. C. C. tortolensis. D–F. Dracolycus larvae spp. D. Dracolycus sp. 1. E–F. Dracolycus sp. 2. Fig. 8. Open in new tabDownload slide Dorsal view of the immature forms of the Puerto Rican bank Leptolycini. Cessator and Dracolycus larvae spp. A–C. Cessator spp. A. C. crypticus. B. C. luquillonis. C. C. tortolensis. D–F. Dracolycus larvae spp. D. Dracolycus sp. 1. E–F. Dracolycus sp. 2. Fig. 9. Open in new tabDownload slide Lateral view of the immature forms of the Puerto Rican bank Leptolycini. Cessator and Dracolycus larvae spp. A–C. Cessator spp. A. C. crypticus. B. C. luquillonis. C–E. Dracolycus larvae spp. C. Dracolycus sp. 1. D–E. Dracolycus sp. 2. Fig. 9. Open in new tabDownload slide Lateral view of the immature forms of the Puerto Rican bank Leptolycini. Cessator and Dracolycus larvae spp. A–C. Cessator spp. A. C. crypticus. B. C. luquillonis. C–E. Dracolycus larvae spp. C. Dracolycus sp. 1. D–E. Dracolycus sp. 2. Fig. 10. Open in new tabDownload slide Ventral view of the immature forms of the Puerto Rican bank Leptolycini. Cessator and Dracolycus larvae spp. A–C. Cessator spp. A. C. crypticus. B. C. luquillonis. C. C. tortolensis. D–F. Dracolycus larvae spp. D. Dracolycus sp. 1. E–F. Dracolycus sp. 2. Fig. 10. Open in new tabDownload slide Ventral view of the immature forms of the Puerto Rican bank Leptolycini. Cessator and Dracolycus larvae spp. A–C. Cessator spp. A. C. crypticus. B. C. luquillonis. C. C. tortolensis. D–F. Dracolycus larvae spp. D. Dracolycus sp. 1. E–F. Dracolycus sp. 2. Fig. 11. Open in new tabDownload slide Distribution map of adult and larvae Cessator in Puerto Rico. A. Distribution of C. crypticus, C. obrienorum, C. luquillonis. B. Distribution of C. crypticus in the El Yunque National Forest. Fig. 11. Open in new tabDownload slide Distribution map of adult and larvae Cessator in Puerto Rico. A. Distribution of C. crypticus, C. obrienorum, C. luquillonis. B. Distribution of C. crypticus in the El Yunque National Forest. Fig. 12. Open in new tabDownload slide Distribution map of adult and larvae Cessator tortolensis and Dracolycus. A. Distribution of C. tortolensis in Puerto Rico and Tortola. B. Distribution of D. chupacabra, D. marshalli and Dracolycus sp 1 and Dracolycus sp 2. Fig. 12. Open in new tabDownload slide Distribution map of adult and larvae Cessator tortolensis and Dracolycus. A. Distribution of C. tortolensis in Puerto Rico and Tortola. B. Distribution of D. chupacabra, D. marshalli and Dracolycus sp 1 and Dracolycus sp 2. CessatorKazantsev, 2009: 93. NanolycusKazantsev, 2013:18 new junior synonym. Differential Diagnosis of Adult Males. Cessator can be distinguished from all other Leptolycini and other Neotropical Lycidae by the combination of the following characters: the tubular, uncompressed antennae (vs. antennae variable in other genera, but to some degree dorsoventrally compressed), with the antennomere III several times longer than the pedicel, which is minute (Figs. 5A–D, 6A–D) (vs. subequal in length in all other Leptolycini; shared feature with Dominopteron), the antennae densely setose, bearing abundant and decumbent setae throughout, with some species having the presence of scaliform setae (Figs. 5A–D, 6A–D) (vs. variable in other genera, but usually densely setose in the antennae of other groups, with bristled setae in Tainopteron and in some of the Leptolycus), the remarkably transverse pronotum (Fig. 5A–D) (vs. longer than wide in Dominopteron, Dracolycus, Electropteron and Leptolycus; the transverse pronotum shared with Tainopteron), the elytra bearing small subcircular cells ranging from strongly developed to weakly developed (vs. smooth elytra in Leptolycus and Dracolycus with even smaller regularly visible cells; strongly developed in Dominopteron and Electropteron) and the short and subtriangular trochanters (vs. long and tubular in Leptolycus; similar shape shared with Dominopteron, Dracolycus, Electropteron and Tainopteron). Type Species. Cessator luquillonisKazantsev, 2009 (by monotypy). Differential Diagnosis of Larvae Cessator larvae can be distinguished from other Leptolycini from the Puerto Rican bank by the following combination: head small, ca. half the length of the pronotum (Fig. 8A–C) (vs. subequal or 2/3 of the size of pronotum in Leptolycus, elongate and narrowed in Dracolycus), the presence of a pair of projections (“little horns”) in the anterior margin of the head, between the nasale (Fig. 8A–C) (vs. projections absent in Leptolycus, weakly developed in Dracolycus), the presence of a distinct and strongly visible body (dorsal and ventral) longitudinal line (Fig. 8A–C) (vs. longitudinal line absent in Leptolycus and Dracolycus) and the presence of long erect setae arising from the tergites and pleurites (Fig. 9A–B) (vs. setae absent in other Leptolycini). Description of Larvae. Body subparallel, dorsoventrally flattened, ranging from creamy, to dark brown or yellow (Figs. 8A–C, 9A–B, 10A–C). Membranes light yellow. Whole body strongly sclerotized, ranging from strongly to weakly punctate (Fig. 8A–C), bearing long erect setae arising from circular cells in the tergites and pleurites (Figs. 8A–C, 9A–B, 10A–C). Dorsal longitudinal line strongly visible and developed throughout (Fig. 8A–C). Head hypognathous, transverse, as wide or narrower than prothorax (Fig. 8A–C), nasale apically blunt (Fig. 8A–C), bearing a pair of strongly developed horn-like projections, one on each side of nasale (Figs. 8A–C, 10A–C). Antennae 2-segmented, inserted in a lateralized anterior portion of head, moderately stalked (Fig. 8A–C); basal antennomere ring-like, very short, up to 6× shorter than antennomere II; apical antennomere apically bulbous. Stemmata present, black, lateralized, positioned anteriad to antennal insertions (Fig. 8A–C). Mandibles symmetrical, ensiform, strongly developed, approximated at base (Fig. 10A–C). Mala transverse. Maxillary palpi slender, 4-segmented, digitiform. Labial palpi minute, seemingly 2-segmented, digitiform, diverging towards apex; prementum entire. Ventral plate medially divided, lateral and anterior angles strongly acute, posteriorly round (Fig. 10A–C). Thorax shape variable: pro-, meso-, and metanotum subtrapezoidal, widest at apex (Figs. 8A–C, 10A–C), lateral margins ranging from round (Fig. 10B) to subparallel (Fig. 10A,C). Prosternum triangular, posteriorly round (Figs. 10A–C). Meso- and metasternum subpentagonal (Fig. 10A–C). Spiracle in mesothorax anteriorly located on top of mesepisternum (Fig. 9A, B). Episterna and epimera strongly developed; episterna widest at base, near pleurites (Fig. 10A–C); epimera slender throughout (Fig. 10A–C). Legs robust, moderately long, tarsungulus distally acute. Abdomen tapering towards apex, slightly telescoped (Fig. 10A–C); sternites wide and transverse (Fig. 10A,C), or subquadrate and medially less sclerotized (Fig. 10B); on side view, with one visible pleurite in abdominal segments I–VIII (Fig. 9A,B); pygopodium whitish, translucid, and weakly developed (Fig. 10A–C). Length: 1.15–4.05 mm. Distribution. Cessator is known to occur in Puerto Rico, Hispaniola, Cuba, and the Virgin Islands. Cessator has species described from Puerto Rico (Cessator luquillonisKazantsev 2009) Cuba (Cessator parda (Zayas 1988)) and from Hispaniola, from Dominican amber (Cessator brodzinskyiFerreira and Ivie 2017) and Cessator anachronicus Ferreira and Ivie 2022 (Ferreira et al. 2022). Several more species are known (but currently undescribed) to occur on Hispaniola and Cuba, and one new species is herein described from the Virgin Islands. Biology. All Leptolycini have been collected in nearly identical situations. Males have been collected in short vegetation by sweeping, in Malaise and Flight Intercept traps, indicating that these are active flying beetles. Larvae have been collected in pitfalls, sifting of leaf litter, Berlese, and pans of Flight Intercept traps, suggesting a high mobility during this life-stage. Taxonomic Notes. After examination of one of the paratypes of Nanolycus gnomusKazantsev 2013 from the Dominican Republic (Kazantsev 2013), we concluded that the species fits all the characteristics of a Cessator mentioned above in the differential diagnosis section. Because of this, we here move Nanolycus gnomusKazantsev, 2013 to Cessator gnomus (Kazantsev, 2013) new combination, rendering NanolycusKazantsev 2013 a new junior synonym of CessatorKazantsev 2009. Cessator luquillonisKazantsev, 2009 Figs. 4, 5A, 6A, 7A, 8B, 9B, 10B, 11A, 11B Cessator luquinollisKazantsev 2009: 93 figs. 3, 32–43; Ferreira and Ivie 2017:58. Examined Type Material. 1: PUERTO RICO: Sierra; de Luquillo 850–900 m; 8-IV-2005; S. Kazantsev leg./ Paratype; Kazantsev det 200 [in red]/ Cessator; luquillonis sp. n.; S. Kazantsev det. 2009/ WIBF051557 (WIBF). Material Examined in Addition to Type Specimens. Adults (n = 19): 1: PUERTO RICO: Naguabo; El Yunque Nat. Forest, S Part; La Sabana recr. are 5.6 km N of; Rio Blanco at rd. PR191; 18º16.1’N, 65º47.6’W, 510 m; 21.vi.-2.vii.2016; Fikáček & Seidel lgt. PR03/ flight intercept trap in a partly in; a sparse rainforest at the river; bank (NMPC). 2: PUERTO RICO: El Verde Sta.; 18.3236ºN,65.8183ºW; 18JUNE-8JULY2017, 1,033 ft; E.L. Spiessberger &; V.S. Ferreira, Malaise #4 (WIBF). 1: PUERTO RICO: El Verde Sta.; 18.3236ºN; 65.8183ºW; 20-23JUNE2017 1,033 ft; V.S. Ferreira &; E.L. Spiessberger, FIT 4/ WIBF051525 (WIBF). 1: PUERTO RICO: El Verde Sta.; 18.3236ºN; 65.8183ºW; 24JUNE2017 1,033 ft; V.S. Ferreira &; E.L. Spiessberger, FIT #4 (WIBF). 1: PUERTO RICO: El Verde Sta.; 18.3218ºN;65.8170ºW; 24JUNE–7JULY2017, 1,225 ft; V.S. Ferreira &; E.L. Spiessberger, FIT 6 (WIBF). 1: PUERTO RICO: 18.3236ºN; 65.8183ºW; 09JULY2017 1,033 ft; Ferreira & Spiessberger; near FIT 4, sweeping (WIBF). 1: PUERTO RICO: El Verde Sta.; 18.3236ºN,65.8183ºW; 273JUNE2017 1,010 ft; V.S. Ferreira &; E.L. Spiessberger, FIT 3/ WIBF051555 (WIBF). 1: PUERTO RICO: El Verde Sta.; 18.3230ºN, 65.8283ºW; 26JUNE2017, 1,117 ft; V.S. Ferreira &; E.L. Spiessberger, FIT 1(WIBF). 1: PUERTO RICO: El Verde Sta.; 18.3218ºN,65.8270ºW; 20JUNE2017, 1,225 ft; V.S. Ferreira &; E.L. Spiessberger, FIT 5 (WIBF). 1: PUERTO RICO: El Verde Sta.; 18.3218ºN,65.8170ºW; 06JULY2017, 1,225 ft; V.S. Ferreira &; E.L. Spiessberger, FIT# 6 (WIBF). 1: PUERTO RICO: Naguabo; El Yunque Nat. Forest, S Part; 4.9 Km N iof Rio Blanco; 18º15.8’N, 65º47.3’W, 495m; 24.vi.-2.vii.2016; Fikáček & Seidel lgt. PR11/ flight intercept trap at the margin; of the rainforest in an area with many flowering Etilingera elatior; plants (NMPC). 1: PUERTO RICO: Ponce, PR-143; B.E. Toro Negro, Mt. Jayuya; 18.1667ºN, 66.5783ºW; 29JUNE2016, 1,265 m; Deler, Fikacek & Seidel/ PR19 Sifting/ WIBF051556 (NMPC). 1: PUERTO RICO: El Verde Sta.; 18.3215ºN, 65.8198ºW; JUNE–JULY2017; Ferreira & Spiessberger (WIBF). 1: PUERTO RICO: El Verde St.; 18.3215ºN,65.8198ºW; 1–4JULY2017, sweeping; Ferreira & Spiessberger/ WIBF041664 (WIBF). 1: PUERTO RICO: Rio Grande Mun.; El Verde Research Station; 18º19.376’N 65º48.997ºW; 17Jul2007 sweeping; coll. M. Gimmel (MLGC). 1: PUERTO RICO: Rio; Grande, El Yunque, Mt.; Britton summit, 541 m; 4–12AUG1999 P.W.; Kovarik, FIT (WIBF). 1: Vista la Sierra; Luquillo Forest; P.R.VIII 9, 1969; H. & A. Howden (CNC). Larvae (n = 9): 1: PUERTO RICO: Arecibo; Bosque Estatal de Cambalache; 18°26.6’N, 66°35.8’W; 70 m 25.vi.2016; Deler, Fikáček & Seidel lgt. PR12/ WIBF 051499 (WIBF). 3: PUERTO RICO: El Verde Field Station; July 2018; T. Sanchez & K. Noa Reyes [WIBF 041694, WIBF 051497, WIBF 051498] (WIBF). 1: PUERTO RICO; El Verde Field Station; ridge tops in Mountain; 15 May–04 Nov 1996; E Nazaro colr; pitfall traps (WIBF). 1: Puerto Rico: Aguas Buenas; Forest at Aguas Buenas; cave 250m, v: 7–17: 1973; leg S. Peck—Ber #; 265. FM (HD) #73-4, Ber:; 93 Kg., 171 L (WIBF). 1: PUERTO RICO: San Germán; Bosque Estatal Maricao, 8.6 km N; of Sabana Grande, 18°8.0’N; 66°59.5’W; 820 m 28.vi.2016; Fikáček & Seidel lgt. PR16 (WIBF). 1: PUERTO RICO: Guayama; W bank of Lago Carite, 9.9 km N; of Guayama; 18°4.4’N, 66°6.3’W; 550 m 23.vi.2016; Deler,Fikáček & Seidel lgt. PR10 [WIBF 051505] (WIBF). 1: PUERTO RICO: Arecibo; Bosque Estatal Río Abajo, 1.3 km SWW of the campsite at Cueva de; las Aguas, 18°19.4’N 66°43.0’W; 330 m, 27.vi.2016 Fikáček & Seidel lgt. PR14C [WIBF 051504] (WIBF). Diagnosis of Adult Males. Cessator luquillonis can be distinguished from all other Cessator from the Puerto Rican bank by having the head in dorsal and lateral view and the pronotum completely black (Fig. 5A) (vs. variable combinations in the other species: head dark brown in Cessator tortolensis and Cessator obrienorum, with the lateral margins in C. tortolensis and C. obrienorum yellow or orange; the pronotum in C. tortolensis, C. obrienorum, and C. crypticus is yellow or orange); the yellow legs and the bicolored elytra (yellow and brown), the latter with the elytra basal 1/6 yellow (Fig. 5A) (vs. legs dark brown in all other species; elytra completely black in C. crypticus and only humeral region of C. tortolensis yellow); the scutellar shield black, with the posterior margin with a shallowly developed, hardly visible median notch (Fig. 5A) (vs. posterior margin of scutellar shield deeply notched in C. tortolensis and C. crypticus, which are both prong-like; yellow, orange or dark-brown in C. obrienorum and C. tortolensis); the gular sutures confluents, directly touching each other (Fig. 6A) (vs. presence of a gular bridge between both sutures in C. tortolensis and C. crypticus); the bullet shaped genitalia, characterized by the median lobe with seemingly absent parameres, 4× shorter than the phallobase, with the presence of a median ventral sickle shaped structured that is apically acuminate in the median lobe (Fig. 7A) (vs. trilobate-like male genitalia in C. tortolensis and fusiform with parameres absent in C. crypticus; similarly shaped to C. obriennorum). Length (pronotum + elytra): 2.27–3.11 mm. Width (across humeri): 0.53–0.72 mm. Diagnosis of Larvae. Cessator luquillonis can be easily separated from the other known Cessator larvae by its coloration dark brown (Figs. 8B, 9B,10B) (vs. yellow in C. crypticus (Figs. 8A, 9A, 10A) and creamy in C. tortolensis), the densely and deeply punctation in the margins of abdominal and thoracic tergites and sternites (Figs. 8B, 9B, 10B) (vs. small and less densely punctuated in C. crypticus (Figs. 8A, 9A, 10A) and smooth in C. tortolensis), the relatively longer and apically spatulate body setation (Figs. 8B, 9B, 10B) (vs. shorter and not apically strongly spatulate in the other species), and by the abdominal sternites subquadrate, medially less sclerotized, leaving a big portion of the membranous part visible in ventral view of abdomen (Figs. 8B, 9B, 10B) (vs. transverse in C. crypticus (Figs. 8A, 9A, 10A), not leaving any membranous part visible in ventral view of abdomen). Length: 4.09–4.61 mm. Distribution. Puerto Rico (Fig. 11). Cessator tortolensis Ferreira and Ivie New Species Figs. 4, 5B, 6B, 7B, 8C, 10C, 12A Zoobank LSID: urn:lsid:zoobank.org:act:772C7359-1CB4-46FB- B46B-1B0806C75BAB Examined Type Material. Adults (n = 14): Holotype: BR: VIRGIN IS: Tortola; Mt.SageNat. Pk. N. side; Mt. Sage, 1,550 ft, steps; 13NOV–10DEC1992, T.R.; Hughes. flt. int. tr. #3 [WIBF002196]. Paratypes: 3: BR: VIRGIN IS: Tortola; Mt.SageNat. Pk. N. side; Mt. Sage, 1,550 ft, steps; 30OCT–13NOV1992, M.A.; Ivie, flt. inter. tr #3 [WIBF002138, WIBF02137, WIBF02139]. 4: BR: VIRGIN IS: Tortola; Mt.SageNat. Pk. N. side; Mt. Sage, 1,550 ft, steps; 23–30OCT1992, M.A. &; L. Ivie, flt. in. tr. #3 [WIBF002117, WIBF002114, WIBF002115, WIBF002116]. 3: BR: VIRGIN IS: Tortola; Mt.SageNat. Pk. N. side; Mt. Sage, 1,550 ft, steps; 13NOV–10DEC1992, T.R.; Hughes. flt. int. tr. #3 [WIBF002197, WIBF002244, WIBF002245]. 1: BR: VIRGIN IS: Tortola; Mt.SageNat. Pk. N. side; Mt. Sage, 1,550 ft, steps; 07–25JUL1994, MIvie &S; Buckling flt. int. tr. #3 [WIBF025536]. 1: BR: VIRGIN IS: Tortola; Mt.SageNat. Pk. S. side; Mt. Sage, 1,520 ft. rest; 14DEC–08JAN1993, M. A.; Ivie flt. inter. tr. #4 [WIBF002586]. 1: BR: VIRGIN IS: Tortola; Mt.SageNat. Pk. N. side; Mt. Sage, 1,550 ft. steps; 10DEC–07JAN1993. M. A.; Ivie flt. inter. tr. #3 [WIBF002257]. Larvae (n = 2): 2: BR: Virgin Is.: Tortola; Mt. Sage Nat. Park; Sage Mt.; 23 OCT 1993; M.A. & L.L. Ivie; mixed litter (WIBF). Material Examined but not Included in the Type Series. Adults (n = 3). 1: PUERTO RICO: El Verde Sta.; 18.3218ºN, 65.8170ºW; 28-29JUNE2017 1,225 ft; V.S. Ferreira &; E.L. Spiessberger, FIT 6/ WIBF051553 (WIBF). 1: PUERTO RICO: El Verde Sta.; 18.3236ºN,65.8183ºW; 27–29JUNE2017 1,010 ft; V.S. Ferreira &; E.L. Spiessberger, FIT 3 (WIBF). 1: PUERTO RICO: El Verde Sta.; 18.3230ºN, 65.8283ºW; 24JUNE2017, 1,117 ft; V.S. Ferreira &; E.L. Spiessberger, FIT 1/ MTEC023637 (WIBF). Etymology. This species has been named after the locality of most of the collected specimens, Tortola, in the British Virgin Islands. Diagnosis of Adult Males. Cessator tortolensis can be easily recognized from all other Cessator from the Puerto Rican bank by having the pronotum distinctly transverse (vs. pronotum not distinctly transverse in the other Cessator), the scutellar shield with a deep notch in the posterior median portion, prong-like (Fig. 5B) (vs. scutellar shield with a shallow notch in the posterior median portion of all other Cessator, not prong-like in Cessator crypticus), the very base of the humeral region yellow (Fig. 5B) (vs. basal 1/6 of elytra with a different color than remaining of the structure in C. luquillonis and C. obrienorum, completely unicolored in C. crypticus) and the gular sutures not directly touching each other, bearing a gular bridge between both sutures (Fig. 6B) (vs. gular sutures confluents, touching each other, in C. luquillonis and C. obrienorum). Description of Male. General coloration: Head, antennae, legs, and elytra brown; pedicel slightly lighter than remaining of antennae (Figs. 5B, 6B), antennomere XI whitish, rarely brown; pronotum, lateral margins of head, scutellar shield, humeral region, and gular region yellow. Body moderately setose. Head: Antennae densely setose, with the presence of scaliform setae (Fig. 5B, 6B). Head strongly alveolate (Fig. 6B); gular sutures not confluents, connected by the presence of a gular bridge between both sutures (Fig. 6B). Thorax: Pronotum strongly transverse, all margins weakly developed, seemingly indistinct, posteriorly flattened (Fig. 5B); scutellar shield with a deep notch in the posterior median portion, prong-like, with very acute apices (Fig. 5B). Elytra slightly dehiscent, strongly ligulate (Fig. 5B); elytral costae weakly visible and developed, with short bristle-like setae throughout elytra (Fig. 5B). Male genitalia trilobate, with phallobase 1/3 longer than parameres; exposed portion of median lobe 1/4 longer than the sum of phallobase and parameres; phallobase posteriorly round; parameres short, 2.5× shorter than exposed portion of median lobe, apices round; median lobe fusiform, apically round, blunt (Fig. 7B). Length (pronotum + elytra): 1.32–1.83 mm. Width (across humeri): 0.36–0.51 mm. Diagnosis of the Larva. Cessator tortolensis is the only known Cessator from the Virgin Islands. The larvae can be readily identified from other Leptolycini larvae by the characteristics presented under the genus description and diagnosis, and from the other known Cessator larvae by its unique creamy color (Figs. 8C, 10C) (vs. dark brown in C. luquillonis and yellow in C. crypticus) and smoother overall texture (Figs. 8C, 10C) (vs. moderately to densely punctuate in the other two species). Length: 1.15 mm. Distribution. British Virgin Islands (Tortola) and Puerto Rico (Fig. 12A). Taxonomic Remarks. Despite minor differences in color pattern in the Puerto Rican specimens (one specimen had the antennae completely brown instead of having the antennomere XI white, and the gular region of other specimen was completely brown instead of yellow), these specimens present an identical external and male genitalia morphology when compared with the specimens from Tortola in the Virgin Islands. Our sampling of specimens from Puerto Rico remains very small, and only two of the three known specimens from the Island could be thoroughly studied (the third specimen has been permanently damaged during the DNA extraction process). Further collection efforts in Puerto Rico using FITs could potentially allow to see more in-depth variations in the Puerto Rican specimens of C. tortolensis and the possible establishment of the identification of more variable characters in a longer series of this species. Cessator obrienorum Ferreira and Ivie New Species Figs. 5C, 6C, 7C Zoobank LSID: urn:lsid:zoobank.org:act:037029A9-B25C-43C1- 938D-51D314DBF8F5 Type Material. Adult male (n = 1): Holotype: 1: PUERTO; RICO 8mi; E Mayaguez; II-9-1969/ Collectors L. &; C.W. O’Brien (WIBF). Etymology. This species is named in honor of Charles W. and Lois B. O’Brien, collectors of the only known specimen of this species. Diagnosis of Adult Males. Cessator obrienorum can be separated from its congenerics from the Puerto Rican bank by possessing the lateral margins of the head, pronotum, scutellar shield orange and 1/6 of the elytra yellow (Fig. 5C) (vs. head and scutellar shield completely black in C. luquillonis and C. crypticus, and only the very base of the humeral region yellow in C. tortolensis), the absence of scaliform setae in the antennae (Fig. 5C, 6C) (vs. presence of scaliform setae in all other Cessator), the gular sutures confluents, directly touching each other (Fig. 6C) (vs. presence of a gular bridge between both sutures in C. tortolensis and C. crypticus, but shared feature with C. luquillonis) and the bullet shaped genitalia, characterized by the median lobe with seemingly absent parameres, 4× longer than the phallobase, with the presence of a median ventral sickle shaped structured that is apically acuminate in the median lobe (shared with C. luquillonis) (Fig. 7C) (vs. trilobate-like in C. tortolensis). Description of Male. General coloration: Dorsal head, antennae (antennomeres I–X) and legs brown; pedicel slightly lighter than remaining of antennae (Figs. 5C, 7C), and antennomere XI white; lateral margins of head, pronotum, scutellar shield, and ventral portion of the head orange (Fig. 5C); elytra bicolored, with basal 1/6 elytra yellow and remaining brown (Fig. 5C). Body densely setose. Head: Antennae densely setose, scaliform setae absent (Figs. 5C, 6C). Head weakly alveolate (Figs. 5C, 6C); gular sutures confluents, directly touching each other (Fig. 6C). Thorax: Pronotum transverse, anterior and posterior margins distinct, side margins weakly developed, posteriorly weakly bisinuate (Fig. 5C); scutellar shield posterior margin with shallow median notch (Fig. 5C). Elytra slightly dehiscent (Fig. 5C); elytral costae distinctly visible, with short decumbent setae throughout the elytra (Fig. 5C). Bullet shaped genitalia, elongate phallobase, and short median lobe; median lobe with seemingly absent parameres, 4× shorter than phallobase, apically in ventral view bearing sickle-shaped structure which are apically acuminate (Fig. 9C). Length (pronotum + elytra): 2.58 mm. Width (across humeri): 0.69 mm. Immature Forms. Unknown. Distribution. Puerto Rico (Fig. 11A). Cessator crypticus New Species Ferreira and Ivie Figs. 4, 5D, 6D, 7D, 8A, 9A, 10A, 11A,B Zoobank LSID: urn:lsid:zoobank.org:act:666921FB-8C46-4B65- 9655-23C4DB92A46E Examined Type Material. Adults (n = 2): Holotype: PUERTO RICO: El Verde Sta.; 18.3236ºN,65.8183ºW; 27–29JUNE2017 1,010 ft; V.S. Ferreira &; E.L. Spiessberger, FIT 3/ WIBF051552 (WIBF). Paratype: PUERTO RICO: El Verde Sta.; 18.3236ºN,65.8183ºW; 30JUNE–1JULY2017 1,010 ft; V.S. Ferreira &; E.L. Spiessberger, FIT 3/ MTEC023638 (WIBF). Examined Material in Addition to the Type Series. Larvae (n = 83): 10: PUERTO RICO: El Verde Field Station; July 2018; T. Sanchez & K. Noa Reyes [WIBF 041695, WIBF 041696, WIBF 051500] (WIBF). 1: PUERTO RICO: El Verde Field Station; 25 June 2017; V. Ferreira &; E.L. Spiessberger [MTEC 023648] (WIBF). 1: PUERTO RICO: Naguabo; El Yunque Nat. Forest, S part; 5.9 km N of Río Blanco above rd.PR191; 18°16.1’N, 65°48.1’W; 575 m, 21.vi.2016; A. Deler-Hernández lgt. PR04/ sifting of leaf litter; accummulations on the slope in; the rainforest area (WIBF). 1: PUERTO RICO: Orocovis; Bosque Estatal Toro Negro, sector Doña Juana; 5.6 km N of Villalba; 18°10.6’N 66°29.7’W 810 m; 29.vi.2016; Deler, Fikáček & Seidel lgt. PR18 (WIBF). 7: PUERTO RICO: Ciales; Bosque Estatal Tres Picachos; 10–10.1 km N of Villalba, around Hacienda Negrón; 18°12.8–12.9’N 66°30.9–31.1’W; 400–550 m 26.vi.2016; Deler, Fikáček & Seidel lgt. PR13b/ sifting of leaf litter in tropical; forest on a slope above river/ sifting of small acummulations of; leaf litter and rotten palm leaves; and fruit in sparse montane; forest at the stream bank (WIBF). 12: PUERTO RICO: Ciales; Bosque Estatal Tres Picachos; 9.8 km N of Villalba, track above Hacienda Negrón; 18°12.7’N 66°31.2’W 670 m 26.vi.2016; Deler, Fikáček & Seidel lgt. PR13a/ sifting of leaf litter in a stony tropical forest on a slope (WIBF). 1: PUERTO RICO: Maricao; For. Res., S. side, 535 m; 7.3 km N Sabana Grande; 26SEP1987, M.A. Ivie; ex forest litter (WIBF). 6: PUERTO RICO; El Verde Research Station; ridge tops in forest; 15 MAY–04 NOV 1996; E. Nazaro colr.; pitfall traps (WIBF). 1: PUERTO RICO; El Verde FS; 23 June 2017; Berlese; 18.323610º N; 065.818310 W; 1,033 ft.; VSF, ELS (WIBF). 43: Puerto Rico: Aguas Buenas; Forest at Aguas Buenas; cave 250 m, v: 7–17: 1973; leg S. Peck—Ber #; 265. FM (HD) #73-4, Ber:; 93 Kg., 171 L (WIBF). Etymology. The species name was given due to the intriguing and cryptic fact that only two adult specimens of this species are known to the authors, even though numerous immatures were found in many localities in Puerto Rico. Diagnosis of Adult Males. Cessator crypticus is readily identifiable from other Cessator by the combination of a head completely black (Figs. 5D, 6D) (vs. bicolored in C. obrienorum and C. tortolensis), a yellow pronotum (Fig. 5D) (vs. black in C. luquillonis; shared with C. obrienorum and C. tortolensis), the unicolored elytra (Fig. 5D) (vs. bicolored in all other Cessator from the Puerto Rican bank), the presence of a gular bridge between both gular sutures (Fig. 6D) (vs. gular sutures confluents, directly touching each other in C. luquillonis and C. obrienorum) and the regular-shaped male genitalia, with parameres absent (Fig. 7D) (vs. bullet shaped in C. luquillonis and C. obrienorum, with apparent reduced parameres). Description of Male. General coloration: Head, antennae, scutellar shield, and legs brown; pedicel slightly lighter than remaining of antennae (Figs. 5D, 6D); pronotum and gular region yellow. Body moderately setose. Head: Antennae densely setose, with the presence of thinner scaliform setae (Fig. 5D, 6D). Head weakly alveolate (Fig. 6D); gular sutures not confluents, connected by presence of gular bridge between both sutures (Fig. 6D). Thorax: Pronotum moderately transverse, anterior, and posterior margins distinct, side margins weakly developed, posteriorly weakly bisinuate (Fig. 5D); scutellar shield posterior margin with moderately deep median notch, prong-like (Fig. 5D). Elytra slightly dehiscent, ligulate (Fig. 5D); elytral costae more visible in basal half, fading towards apex, with short bristle-like setae throughout elytra (Fig. 5D). Male genitalia elongate, fusiform; phallobase 1/3 longer than median lobe, parameres absent; median lobe apically round, blunt (Fig. 7D). Length (pronotum + elytra): 2 mm. Width (across humeri): 0.59 mm. Diagnosis of the Larva. See comparative diagnosis under C. luquillonis. Length: 1.56–4.05 mm. Distribution. Puerto Rico (Fig. 11A and B). A Key to the Adult Male Cessator of the Puerto Rican Bank 1. Pronotum and head when viewed dorsally black (Fig. 5A); legs yellow (Fig. 5A) Cessator luquillonisKazantsev 2009 1.’ Pronotum and head with a different color combination; legs brown 2 2. Elytra unicolored, dark brown (Fig. 5D) Cessator crypticus new species 2.’ Elytra bicolored, sometimes only the base of the humeri region have a different color 3 3. Scutellar shield with a deep notch in the posterior median portion, prong-like; pronotum remarkably transverse (Fig. 5B); gular sutures not directly touching each other, with the presence of a gular bridge between both sutures (Fig. 6B) Cessator tortolensisnew species 3.’ Scutellar shield with a shallow notch in the posterior median portion (Fig. 5C); the gular sutures confluents, directly touching each other (Fig. 6C) Cessator obrienorumnew species Dracolycus Ferreira and Ivie New Genus Figs. 5E–F, 6E–F, 7E–F, 8D–F, 9C–E, 10D–F, 12B Zoobank LSID: urn:lsid:zoobank.org:act:0E728457-BEAF-406B- A3F5-5E962F1E8444 Differential Diagnosis of Adult Males. Dracolycus can be readily identified from all other Leptolycini by the combination of the following characters: the filiform, dorsoventrally compressed antennae (Fig. 5E–F) (vs. variable in other Leptolycini: tubular and not compressed in Cessator; flabellate, filiform and subserrate and strongly compressed in Leptolycus; filiform and moderately compressed in Tainopteron), with relatively wide antennomeres IV–XI (Fig. 5E–F) (vs. narrowed in most Leptolycini), bearing short decumbent setae (vs. variable across the Leptolycini: with abundant decumbent setae in Cessator, Dominopteron, and some Leptolycus; bearing long erect bristle-like setae in some Leptolycus and Tainopteron), the absence of scaliform setae (vs. presence of scaliform setae in some Leptolycus and Cessator), and the pedicel and antennomere III subequal in size (Fig. 5E–F) (vs. pedicel much smaller than antennomere III in Dominopteron and Cessator). Dracolycus also possesses a pronotum that is nearly as long as wide, slightly trapezoidal, with strongly developed margins (Fig. 5E–F) (vs. pronotum shaped variable in other Leptolycini: as long as wide in most Leptolycus, with overall indistinct margins in this genus; as long as wide in Dominopteron, with strongly distinct margins; transverse and either imarginate or weakly developed in Cessator and Tainopteron), and the presence of a strongly visible median longitudinal carina (Fig. 5E–F) (vs. no visible carina in other Leptolycini). The new genus is also unique among the other Leptolycini by possessing a cordiform scutellar shield (Fig. 5E–F) (vs. subquadrate or rectangular in other Leptolycini), a developed elytra, covering or surpassing the abdomen tip (Fig. 5E–F) (vs. weakly developed in most genera, normally strongly dehiscent, sometimes ligulate in other Leptolycini, usually leaving part of the abdomen visible), smooth textured, with no apparent cells or reticulations other than the two longitudinal costae in each elytron (Fig. 5E–F) (vs. variable across the other Leptolycini, but always presenting cells or reticulations), and a sword-like genitalia, with the exposed portion of the median lobe twice the length of parameres and phallobase (Fig. 7E–F) (vs. highly variable in other Leptolycini, but not as described for Dracolycus). Etymology. The name is an allusion to the author’s hypothesis that the adult females/larvae of this new taxon (Dracolycus sp. 1 and sp 2) resembles the shape of a dragon or a serpent, in particular the ones depicted in popular Japanese culture. Therefore, Draco-, from Ancient Greek δράκων (drákōn), meaning serpent, dragon, and Lycus, in allusion to the type genus of the family Lycidae. Gender masculine. Type Species. Dracolycus chupacabra Ferreira and Ivie. Description of Adult Males. General dorsal coloration and pubescence ranging from brown to yellow, antennomere XI yellow (Fig. 5E–F). Body densely setose, with short, decumbent setation throughout. Head: as long as wide, widest at eyes, posteriorly partially covered by pronotum, hypognathous, frons strongly, or weakly bulgy (Fig. 6E–F). Eyes hemispherical, projecting anterolaterally in dorsal view, coarsely granulate (Fig. 6E–F). Mouthparts: Maxillary palp four-segmented, bearing setae as long as palp; terminal palpomeres acuminate. Labial palps and mandibles seemingly absent. Posterior margin of epistoma clearly emarginate; labrum distinct from clypeus, transverse and small, densely setose. Gular sutures confluents, directly touching each other, gular sutural bridge absent (Fig. 6E–F). Antennae: filiform, densely setose, dorsoventrally flattened from IV–XI, inserted in gibbous prominence at anterior distal portion of head, 11-segmented; in resting position reaching basal third or half of elytra (Fig. 5E–F); scape pyriform, approximated at base, subconical; pedicel and antennomere III short, subequal in size and length, pedicel pyriform, shortened; antennomere 4–5x longer than precedent; antennomere V 4/5 of size of precedent; antennomeres V–X subequal in size and shape, slightly decreasing in length towards apex, XI round apically (Fig. 5E–F). Thorax: Prothorax: Pronotum as long as wide, trapezoidal (Fig. 5E–F); margins distinct and developed, side margins slightly arcuate, posteriorly subtly sinuated; angles round, posteriorly slightly acute; pronotum with a longitudinal median carina strongly visible; inner area adjacent to side margins slightly concave (Fig. 5E–F). Hypomeron slightly concave. Prosternum Y-shaped, anteriorly developed extending over gular region or weakly developed (Fig. 6E–F); posteriorly strongly or moderately acuminate. Mesothorax: Mesospiracles not visible in the examined specimens. Mesoventrite trapezoidal. Scutellar shield cordiform, protruded (Fig. 5E–F). Metathorax: Metaventrite convex, posterolateral angles acute; metadiscrimen complete. Elytra: Ranging from slightly to distinctly dehiscent (Fig. 5E–F), strongly developed, slightly ligulate or expanded apically, ca. 5× longer than pronotum; smooth textured, with small, seemingly absent circular weakly developed cells (Fig. 5E–F); bearing two elytral costae fused apically; secondary costae absent (Fig. 5E–F). Legs: slender, elongate, dorsoventrally strongly flattened; protrochanthins slender and exposed (Fig. 5E–F); pro- and mesocoxae conical, obliquely positioned; procoxae contiguous, metacoxae transverse, with posterior face weakly excavated. Trochanters subtriangular, not tubular. Femora parallel sided (Fig. 5E–F). Tibiae clavate, slender in basal half, elongate; tibiae slightly longer than femora or subequal in length. Tarsal formula 5-5-5, all tarsomeres narrowed. Abdomen of males with eight visible ventrites; ventrite VII medially apically notched; ventrite VIII elongate, blade like, apically acuminate. Male genitalia typical elateriformia: trilobate, symmetrical (Fig. 7E–F); phallobase posteriorly round, median suture absent (Fig. 7E–F); parameres short, apically round, co-adapted with median lobe; median lobe elongate, exposed portion ranging from 2–2.5x the length of parameres + phallobase, apex acuminate or round (Fig. 7E–F). Distribution. Specimens are known from Puerto Rico, from the El Verde Field Station (see descriptions below) (Fig. 12B). One unnamed fossil specimen (apparently the same species as Dracolycus marshalli) is known from Dominican Amber, depicted in Wu (1997:179). Despite several attempts of contacting Wu regarding the amber specimen location (listed in his original publication as belonging to his private collection), we were not able to locate and communicate with the author about his specimen. Biology. The only known male specimens of this genus were collected using Malaise traps. The specimens tentatively associated with the immature forms of Dracolycus have been collected by sifting, Winkler and Berlese in moist leaf litter and in flight intercept traps (FIT). Larvae Tentatively Associated with Dracolycus. Based in our extensive sampling of Leptolycini specimens from the Puerto Rican bank, we were able to provide direct and indirect associations of the immatures and their correspondent adults of two out of four genera known (Cessator and Leptolycus) for the region, providing the first descriptions and diagnosis of these beetles (see association DNA and biogeography-based associations in the beginning of the results section and diagnosis and descriptions under each specific genus). To this moment, a definitive association between adults and immature forms of Dracolycus and Tainopteron remains to be provided. Based in our study of the immature forms of Cessator and Leptolycus, it is reasonable to extrapolate that within the Leptolycini, their immature and respective adult forms possess an overall similar size (length and width). The larvae that we are provisionally treating as Dracolycus are relatively large specimens, with an elongate and slender body, that have roughly the same size expected of the potential immature forms of Dracolycus, which are the largest known Leptolycini. On the other hand, Tainopteron are ca. 10× smaller than Dracolycus, possessing a compact body type and miniaturized overall structures, and we predict that the correspondent larva for the genus will have a more or less similar size that its adult. We are very confident that the larvae that we are tentatively treating as the immature form of Dracolycus is a lycid in the tribe Leptolycini, which is supported by their unmistakable similarity with the herein described larvae of other genera in this tribe. However, knowing of the limitations of our logical inference, and since we were not able to provide any direct association of these specimens with their adult forms, we decided to briefly diagnose the two different larva types found without providing any specific names for them, with the goal of exposing and highlighting the existence of this type of larva in the formal published Lycidae literature. In addition to providing high-quality photographs and brief diagnosis of these larval type, we provided and generated COI sequences (MTEC 023640, WIBF 041511, WIBF 051523), with the hopes that in the near future a definitive test of our hypotheses can be provided, and DNA of adult males can be matched with these larvae. The diagnosis and brief descriptions for these larval types are provided after the descriptions of the two adult males of Dracolycus. Dracolycus chupacabra Ferreira and Ivie New Species Figs. 5E, 6E, 7E, 12B Zoobank LSID: urn:lsid:zoobank.org:act:A92955C1-8754-4E8E- B5EE-6186C7DE705F Examined Type Material. Adults (n = 1): Holotype: 1: PUERTO RICO: El Verde Sta.; 18.3236ºN; 65.8183ºW; 18JUNE–8JULY2017, 1,033 ft; E.L. Spiessberger &; V.S. Ferreira Malaise#4/ WIBF041538 (WIBF). Etymology. The species is named after the famous Chupacabra (“goat-sucker”) urban legend, which originated in Puerto Rico. The real Chupacabra might be similar to the species herein described. Diagnosis of Adult Males. Dracolycus chupacabra can be easily separated from Dracolycus marshalli by its brown pronotum (Fig. 5E) (vs. pronotum yellow in D. marshalli), the strongly dehiscent and more slender elytra (Fig. 5E) (vs. elytra weakly dehiscent, expanded apically in D. marshalli), the fusion of the elytral costae similar throughout, not expanded apically (Fig. 5E) (vs. elytra costae fusion expanded apically in D. marshalli) and by the apex of the median lobe of the male genitalia lanceolate (Fig. 7E) (vs. apex of median lobe round in D. marshalli). Description of Male. General coloration: Head, antennae, pronotum and elytra brown (Fig. 5E); scutellar shield, antennomere XI, coxae, trochanters, apical third of pro- and mesofemora and metatibiae (with exception of middle third which is brown) and abdomen yellow. Head: frons weakly developed, not bulgy (Fig. 6E). Maxillary palps approximated at base, terminal maxillary palpomere distinctly acuminate. Prosternum anteriorly developed, extending over gular region (Fig. 5E); posteriorly moderately acuminate (Fig. 5E). Metanepisternum crescent shaped throughout. Elytra dehiscent, ligulate (Fig. 5E); fusion of the elytral costae similar throughout, not expanded apically (Fig. 5E). Male genitalia: median lobe lanceolate, apically acuminate (Fig. 7E); exposed portion of median lobe 2× longer than phallobase + parameres (Fig. 7E). Length (pronotum + elytra): 4.88 mm. Width (across humeri): 0.92 mm. Distribution. Puerto Rico, El Verde Field Station (Fig. 12B). Dracolycus marshalli Ferreira and Ivie New Species Figs. 5F, 6F, 7F, 12B Zoobank LSID: urn:lsid:zoobank.org:act:F578DF4C-67FA-4859- B349-2CD3C4905E4D Type Material. Adults (n = 1): Holotype: 1: P.R.: El Verde; malaise trap; in forest; 1–2. ii. 1989; S.A. Marshall/ WIBF051558 (DEBU). Etymology. This species is named in honor of Steve A. Marshall, collector of this species. Diagnosis of Adult Males. See diagnosis under Dracolycus chupacabra. Description of Male. General coloration: Head, antennae, elytra, antennae and legs (with exception of the very apex of femora and base of tibiae yellow) brown (Fig. 5F); pronotum, scutellar shield, antennomere XI, coxae, trochanters, thorax and middle region of ventrites yellow. Head: frons strongly bulgy. Maxillary palps separated at base, palpomeres terminal maxillary palpomere acuminate, apex not strongly acute (Fig. 6F). Prosternum anteriorly weakly developed, posteriorly strongly acuminate. Metanepisternum crescent shaped, basal half anteriorly blunt, with outer margin in ventral view abruptly acute, strongly acuminate. Elytra only slightly dehiscent, expanding towards apex (Fig. 5F); fusion of elytral costae expanding towards apex, widest in elytral 3/4 (Fig. 5F). Male genitalia: median lobe lanceolate, apex round; exposed portion of median lobe 2.5× longer than phallobase + parameres (Fig. 7F). Length (pronotum + elytra): 6.64 mm. Width (across humeri): 1.12 mm. Distribution. Puerto Rico, El Verde Field Station (Fig. 12B). A Key to the Adult Males of Dracolycus of the Puerto Rican Bank 1. Pronotum and elytra dark brown; fusion of elytral costae similar throughout (Fig. 5E); elytra strongly dehiscent, slendered, not expanded apically (Fig. 5E)… Dracolycus chupacabra Ferreira and Ivie new species 1.’ Pronotum yellow, elytra dark brown (Fig. 5F); fusion of elytral costae distinctly expanded in mid elytra; elytra weakly dehiscent, expanded apically (Fig. 5F)… Dracolycus marshalli Ferreira and Ivie new species Larvae Tentatively Associated with Dracolycus Figs. 4, 8D–F, 9C–E, 10D–F, 12B Description of Larvae. Body parallel-sided, dorsoventrally flattened, ranging from pale yellow to dark brown (Figs. 8D–F, 9C–E, 10D–F), legs pale yellow (Fig. 8D–F). Membranes color matching overall color of the body. Body strongly sclerotized and compact, dorsally presenting two longitudinal median parallel rows of punctations throughout (Figs. 8D–F, 9C–E, 10D–F), punctations depth and arrangement variable in the different species; presence of short, sparsely distributed trichoid setation in tergites (Figs. 8D–F, 9C–E). Head hypognathous, transverse, posteriorly as wide as prothorax (Fig. 8D–F), anteriorly strongly wide (Fig. 8D–F); nasale not angulate, bearing a pair of weakly developed and short setae-like projections, one on each side of nasale (Fig. 8D–F). Antennae 2-segmented, inserted in a lateralized anterior portion of head, strongly stalked (Fig. 8D–F); antennomere I encapsulating basal half of second; antennomere II apically round, plug-like. Stemmata present, black, lateralized, positioned anteriad to antennal insertions (Fig. 8D–F). Mandibles symmetrical, ensiform, weakly to moderately developed, approximated at base (Fig. 9C–E). Mala transverse. Maxillary palpi slender, digitiform, 4-segmented. Labial palpi minute, seemingly 2-segmented; prementum entire. Ventral plate medially divided, lateral and anterior angles strongly acute, posteriorly medially angulate, round or shallowly notched. Thorax distinctly longer than wide, elongate and strongly narrowed (Fig. 8D–F), marginate, angles round, subparallel (Fig. 9C–E). Prosternum triangular, elongate, apically widest (Fig. 10C–E), anterior angles moderately acute, posterior angle round, blunt or acute (Fig. 10D–F). Meso- and metasternum heptagonal, elongate, anterior and posterior angles acute (Fig. 10D–F). Spiracle in mesothorax anteriorly located on top of mesepisternum. Episterna and epimera strongly developed; episterna shorter, half the size of epimeron, widest at base (Fig. 10D–F); epimera slender throughout (Fig. 10D–F). Legs robust, moderately long, tarsungulus distally acute (Fig. 10D–F). Abdomen tapering towards apex (Fig. 10D–F); sternites elongate, longer than wide and transverse. Spiracle plate (upper pleurite) strongly developed and elongate (Figs. 9C–E, 10D–F); lower pleurite strongly reduced, lobed, a third of the length and half the width of the upper, on side view located in posterior ventral portion of upper pleurite, in ventral view of abdomen, located posterolaterad of abdominal sternites (Fig. 10D–F). Pygopodium round, whitish and translucid, weakly developed (Fig. 10D–F). Fig. 13. Open in new tabDownload slide Male adult habiti of the Puerto Rican bank Leptolycini—Leptolycus. A. Leptolycus flavicollis. B. Leptolycus viensis. C. Leptolycus falsoheterocornis new species. D. Leptolycus puellus. E-G. Leptolycus heterocornis species complex. E. Color morph 1. F. Color morph 2. G. Color morph 3. H. Color morph 4. Fig. 13. Open in new tabDownload slide Male adult habiti of the Puerto Rican bank Leptolycini—Leptolycus. A. Leptolycus flavicollis. B. Leptolycus viensis. C. Leptolycus falsoheterocornis new species. D. Leptolycus puellus. E-G. Leptolycus heterocornis species complex. E. Color morph 1. F. Color morph 2. G. Color morph 3. H. Color morph 4. Length: 3.44–4.21 mm. Dracolycus sp. 1 Figs. 8D, 9C, 10D Examined larvae(n = 22). 1: PUERTO RICO: El Verde Field Station; Mixed forest, 500 m; M500G; July 10, 2017; J. Nelsen and N. Scavo colrs./ M500 g/ July 10; 17 (WIBF). 1: PUERTO RICO: El Verde Field Station; Mixed forest, 1,000 m; MFT1000G; June 26, 2017; J. Nelsen and N. Scavo colrs./ 06.26.2017/ MFT 1000G [MTEC 023640] (WIBF). 1: PUERTO RICO: El Verde Sta.; 18.3236ºN,65.8183ºW; 1JUL2017 1,010 ft; V.S. Ferreira &; E.L. Spiessberger, FIT 3 [MTEC 023642] (WIBF). 1: PUERTO RICO: Guayama; W bank of Lago Carite, 9.9 km N; of Guayama; 18°4.4’N, 66°6.3’W; 550 m; 23.vi.2016; Deler,Fikáček & Seidel lgt. PR10 [WIBF 051523] (WIBF). 4: PUERTO RICO: San Germán; Bosque Estatal Maricao, 8.6 km N of Sabana Grande; 18°8.0’N 66°59.5’W; 820 m 28.vi.2016; Fikáček & Seidel lgt. PR16 (WIBF). 14: PUERTO RICO; El Verde Research Station; ridge tops in forest; 15 MAY–04 NOV 1996; E. Nazaro colr.; pitfall traps (WIBF). Differential Diagnosis. Dracolycus larva sp 1 can be separated from Dracolycus larva sp 2 by the antennal insertion projections not strongly stalked (Figs. 8D, 9C) (vs. strongly stalked in Dracolycus sp. 2 (Figs. 8E–F, 9D–E)), the two dorsal longitudinal parallel rows of punctations distinctly strongly impressed and demarcated throughout (Fig. 8D) (vs. not strongly impressed in Dracolycus sp. 2 (Figs. 8E–F, 9D–E)), which are distinctly connected apically by a transversal row in the thorax (Fig. 8D) (vs. not distinctly connected in Dracolycus sp. 2 (Figs. 8E–F)), by possessing the apex of the posterior portion of ventral plate of the head medially acuminate and round (Fig. 10D) (vs. posterior portion of ventral plate of the head notched apically in Dracolycus sp. 2 (Figs. 10E–F)), and by the yellow coloration (Fig. 10D) (vs. pale yellow or brown in Dracolycus sp. 2 (Figs. 10E–F)). Length: 3.7 mm. Dracolycus sp. 2 Figs. 8E–F, 9D–E, 10E–F Yellow Larvae(n = 10). 1: PUERTO RICO: Arecibo; Bosque Estatal Río Abajo, 1.3 km SWW of the campsite at Cueva de las Aguas, 18°19.4’N; 66°43.0’W; 330 m, 27.vi.2016; Fikáček & Seidel lgt./ sifting of leaf litter and mosses; on trunks and stones in sparse; lowland forest on limestone bedrock/ side pools of a stony stream with many decaying leaves (WIBF). 1: M400J [WIBF 041511] (WIBF). 1: PUERTO RICO: El Verde Sta.; 18.3230ºN,65.8183ºW; JULY 2017, 1,010 ft; V.S. Ferreira &; E.L. Spiessberger, FIT 3 [MTEC 023641] (WIBF). 1: Puerto Rico, El Verde Field Station; 22 Jun 2017; leaf litter; VSF, ELS [WIBF 051502] (WIBF). 1: Puerto Rico: Aguas Buenas; Forest at Aguas Buenas; cave 250 m, v: 7-17: 1973; leg S. Peck—Ber #; 265. FM (HD) #73-4, Ber:; 93 Kg., 171 L (WIBF). 5: PUERTO RICO; El Verde Research Station; ridge tops in forest; 15 MAY–04 NOV 1996; E. Nazaro colr.; pitfall traps (WIBF). Grey Larvae (n = 4). 1: PUERTO RICO: Guayama; W bank of Lago Carite, 9.9 km N; of Guayama 18°4.4’N, 66°6.3’W; 550 m; 23.vi.2016; Deler, Fikáček & Seidel lgt. PR10 [WIBF 051522] (WIBF). 1: PUERTO RICO: Ciales; Bosque Estatal Tres Picachos; 10–10.1 km N of Villalba, around Hacienda Negrón; 18°12.8–12.9’N 66°30.9–31.1’W; 400–550 m; 26.vi.2016; Deler, Fikáček & Seidel lgt. PR13b (WIBF). 1: PUERTO RICO: Río Grande; El Yunque Nat. Forest, N part; 9.45 km SE of Río Grande at rd. PR-191; 18°18.6’N, 65°46.7’W; 595 m, 22.vi.2016; Deler, Fikáček & Seidel PR07/ sifting of small accummulations; of moist leaf litter in a high; sparse rainforest at the bank of; a small stony river [WIBF 051501] (WIBF). 1: PUERTO RICO: El Verde Field Station; July 2018; T. Sanchez & K. Noa Reyes (WIBF). Differential Diagnosis. See diagnosis under Dracolycus sp. 1. Length: 3.44–4.21 mm. LeptolycusLeng and Mutchler 1922 Figs. 4, 13A–H, 14A-H, 7H–N, 15A–G, 16A–G, 17A–G, 18A–E, 19A, 20 Fig. 14. Open in new tabDownload slide Head ventral detail of the adult male Puerto Rican bank Leptolycini—Leptolycus. A. L. flavicollis. B. L. viensis new species. C. L. falsoheterocornis new species. D. L. puellus. E–H. L. heterocornis color morphs. Fig. 14. Open in new tabDownload slide Head ventral detail of the adult male Puerto Rican bank Leptolycini—Leptolycus. A. L. flavicollis. B. L. viensis new species. C. L. falsoheterocornis new species. D. L. puellus. E–H. L. heterocornis color morphs. Fig. 15. Open in new tabDownload slide Dorsal view of the immature forms of the Puerto Rican bank Leptolycini. Leptolycus larvae spp. A–C. L. heterocornis color morphs. D. Leptolycus sp. 1. E. Leptolycus sp. 2. F. Leptolycus sp. 3. G. L. viensis. Fig. 15. Open in new tabDownload slide Dorsal view of the immature forms of the Puerto Rican bank Leptolycini. Leptolycus larvae spp. A–C. L. heterocornis color morphs. D. Leptolycus sp. 1. E. Leptolycus sp. 2. F. Leptolycus sp. 3. G. L. viensis. Fig. 16. Open in new tabDownload slide Lateral view of the immature forms of the Puerto Rican bank Leptolycini. Leptolycus larvae spp. A–C. L. heterocornis color morphs. D. Leptolycus sp. 1. E. Leptolycus sp. 2. F. Leptolycus sp. 3. G. L. viensis. Fig. 16. Open in new tabDownload slide Lateral view of the immature forms of the Puerto Rican bank Leptolycini. Leptolycus larvae spp. A–C. L. heterocornis color morphs. D. Leptolycus sp. 1. E. Leptolycus sp. 2. F. Leptolycus sp. 3. G. L. viensis. Fig. 17. Open in new tabDownload slide Ventral view of the immature forms of the Puerto Rican bank Leptolycini. Leptolycus larvae spp. A–C. L. heterocornis color morphs. D. Leptolycus sp. 1. E. Leptolycus sp. 2. F. Leptolycus sp. 3. G. L. viensis. Fig. 17. Open in new tabDownload slide Ventral view of the immature forms of the Puerto Rican bank Leptolycini. Leptolycus larvae spp. A–C. L. heterocornis color morphs. D. Leptolycus sp. 1. E. Leptolycus sp. 2. F. Leptolycus sp. 3. G. L. viensis. Fig. 18. Open in new tabDownload slide Extreme paedomorphic adult female of L. viensis. A. Dorsal habitus. B. Lateral habitus. C. Ventral habitus. D. Head detail, dorsal view. E. Head detail, ventral view. Fig. 18. Open in new tabDownload slide Extreme paedomorphic adult female of L. viensis. A. Dorsal habitus. B. Lateral habitus. C. Ventral habitus. D. Head detail, dorsal view. E. Head detail, ventral view. Fig. 19. Open in new tabDownload slide Distribution map of adult Leptolycus and Tainopteron species in Puerto Rico. A. Distribution of L. puellus, L. flavicollis, L. falsoheterocornis. B. Distribution of Tainopteron milleri. Fig. 19. Open in new tabDownload slide Distribution map of adult Leptolycus and Tainopteron species in Puerto Rico. A. Distribution of L. puellus, L. flavicollis, L. falsoheterocornis. B. Distribution of Tainopteron milleri. Fig. 20. Open in new tabDownload slide Distribution map of adult Leptolycus heterocornis and Leptolycus viensis. A. Leptolycus heterocornis (adult males and larvae morphs) distribution in Puerto Rico. B. Leptolycus viensis (adults males and females, larvae) distribution in the Virgin Island. Fig. 20. Open in new tabDownload slide Distribution map of adult Leptolycus heterocornis and Leptolycus viensis. A. Leptolycus heterocornis (adult males and larvae morphs) distribution in Puerto Rico. B. Leptolycus viensis (adults males and females, larvae) distribution in the Virgin Island. LeptolycusLeng and Mutchler, 1922: 430; Kleine 1933: 18; Beatty 1944: 136; Blackwelder 1945: 343; Bocak and Bocakova 1990: 639; Bocak 2001: 204; Bocak and Bocakova 2008: 714; Kazantsev 2009: 88, 91. Leptolycus (Baholycus) Bocak, 2001: 206; Kazantsev 2009: 95. New junior synonym. Type Species. Leptolycus heterocornisLeng and Mutchler, 1922 (by monotypy). Differential Diagnosis of Adult Males. Leptolycus is one of the most easily recognizable Lycidae in the New World. The genus can be easily recognized among other Lycidae by the following combination of characters: the head longer than wide, widest at middle where the eyes are positioned (Fig. 13A–H) (vs. variable in other Leptolycini, but not as distinct as in Leptolycus), the pedicel and antennomere III subequal in size (Fig. 13A–H) (vs. pedicel much smaller than antennomere III in Dominopteron and Cessator) and the pronotum as long as wide, distinctly constricted medially, with side margins indistinct, seemingly absent (Fig. 13A–H) (vs. pronotum shaped variable in other Leptolycini: as long as wide in Dracolycus and Dominopteron, with margins distinctly visible and not constricted medially; transverse and either imarginate or weakly developed in Cessator and Tainopteron). The antennae in Leptolycus are quite variable in the different species (Fig. 13A–H), surpassing mid-elytra when in resting position in most species, ranging from filiform (in L. heterocornis with the inner margins irregularly shaped, almost crenulate), subserrate (in L. puellus) or flabellate, possessing either short, decumbent setae, bristle-like setae, or a combination of short decumbent setae and scaliform cells. Leptolycus also have the elytra remarkably dehiscent and ligulate, tapered towards apex, always exposing part of the abdomen and the membranous wings underneath (Fig. 13A–H) (vs. variable in the other Leptolycini, usually not strongly dehiscent, exposed portions of abdomen and wings not as in Leptolycus), the elytra with a smooth texture, with cells weakly developed or seemingly absent (Fig. 13A–H) (vs. strongly developed cells in Dominopteron and Electropteron), and very long and slender legs with the metatrochanters distinctly elongate and tubular (vs. variable in other Leptolycini, metatrochanters never tubular nor elongate). The male genitalia is strongly simplified and highly similar throughout the different species in the genus, characterized by a lanceolate median lobe, apically acuminate, and by the absence of parameres (Fig. 7H–N), with the median lobe 0.5–2× the length of phallobase (vs. variable in other Leptolycini: regular trilobate shape in Dominopteron, Tainopteron and some Cessator; C. crypticus has absent parameres whilst C. obrienorum and C. luquillonis have seemingly absent parameres). Differential Diagnosis of Larvae. Leptolycus larvae can be distinguished from the other two Leptolycini genera from the Puerto Rican bank by the long and wide head, subequal or 2/3 the size of the pronotum (Fig. 16A–G) (vs. head ca. half the size of the pronotum in Cessator and strongly narrow in Dracolycus), the absence of a median longitudinal line in dorsal and ventral view (Figs. 15A–G, 17A–G) (vs. presence of a median longitudinal line in Cessator), the strongly alveolate and glabrous body (Figs. 15A–G, 16A–G, 17A–G) (vs. non alveolate body in Cessator and Dracolycus; body possessing long erect setae in Cessator) and the subpentagonal shape of the meso- and metasternum (Fig. 17A–G) (vs. subcircular and elongate in Cessator, kite-like, heptagonal in Dracolycus). Description of Larvae. Body subparallel, dorsoventrally flattened, ranging from creamy, to black or dark brown (Fig. 15A–G); metathorax entirely or partially yellow or black; tergites IV-VI variable in color, entire black, creamy, or partially yellow (Fig. 15A–G). Membranes light brown. Whole body weakly sclerotized, strongly alveolate (Fig. 15A–G). Head hypognathous, transverse, as wide or wider than prothorax, nasale apically bisinuate, or straight (Fig. 15A–G). Antennae 2-segmented, inserted in lateralized anterior portion of head, moderately to distinctly stalked (Fig. 15A–G); basal antennomere ring-like, very short, up to 6× shorter than antennomere II; apical antennomere apically bulbous. Eyes present, black, lateralized, positioned anteriad to antennal insertions (Fig. 15A–G). Mandibles symmetrical, falciform, inner margin sharpened, ranging from small to strongly developed (Fig. 16A–G), approximated at base. Mala transverse. Maxillary palpi slender, digitiform, 4-segmented. Labial palpi minute, seemingly 2-segmented or absent; prementum entire, not medially divided. Ventral plate entire, lateral and anterior angles strongly to moderately acute, posteriorly round (Fig. 16A–G). Thorax shape variable: pro-, meso-, and metanotum ranging from square to rectangular, moderately to distinctly transverse (Fig. 15A–G). Prosternum triangular, posteriorly round (Fig. 17A–G). Meso- and metasternum subpentagonal (Fig. 17A–G). Spiracle in mesothorax anteriorly located on top of mesepisternum (Fig. 16A–G). Episterna and epimera strongly developed; episterna widest at base, near pleura, epimera slender throughout (Fig. 8A–F). Legs robust, moderately long, tarsungulus distally acute (Fig. 17A–G). Abdomen tapering towards apex, slightly telescoped (Figs. 16A–F, 17A–G), with paired lateral pleurites in abdominal segments I–VIII (Figs. 16A–G, 17A–G), pygopodium semicircular, whitish, translucid, strongly developed (Figs. 16A–G, 17A–G). Differential Diagnosis of Extreme Paedomorphic Female. See diagnosis under Leptolycus viensis. Distribution. Cuba, Hispaniola, and the Puerto Rican Bank (Figs. 19A, 20A,B). Even though Beatty (1944:136) and Miskimen and Bond (1970:83) indicated the presence of Leptolycus in St. Croix, no vouchers of those specimens were ever found, nor were any other specimens collected from there by MAI and collaborators despite repeated expeditions to the island. Such patterns of incorrect distribution from West Indies beetles is not unheard of, especially for Lycidae and Lampyridae (see the emblematic case of Thonalmus bicolor (Linnaeus, 1763), endemic to Hispaniola but recorded as occurring in several WI islands and Africa in Ferreira and Ivie (pers. comm.) and other cases in lampyrids in Keller and Branham 2021). In addition to this, MAI met George Miskimen at the University of Puerto Rico at Mayaguez and he admitted that some of the beetle records that he included in his study were just things he thought should be there, and not that he ever saw them. Taking these in consideration, and with the absence of vouchers and no further evidence of specimens of Leptolycus from this island, we consider this to be an incorrect distribution record. Biology. Similar to Cessator and Dracolycus, males have been collected in short vegetation by sweeping and in Malaise and Flight Intercept traps, indicating that these are active flying beetles. Larvae and adult females were collected in pitfalls, sifting on leaf litter, Berlese, and pans of Flight Intercept traps, suggesting a high mobility in this life-stage. Note on the Interspecific Variations of Leptolycus. After the examination and study of several type specimens of Hispaniolan species placed in Baholycus (a revision of Leptolycus is in preparation by the authors) and the examination of the paratype of L. puellus (Puerto Rico), we concluded that all these taxa fit the characteristics mentioned above for Leptolycus strictu sensu. The only encountered variations were in the shape of the antennae (mentioned as flabellate in Baholycus and L. puellus, but truly subserrate in the latter) and the presence of scaliform setae and the mammiform tubercles in the pronotum of some of the Hispaniolan Baholycus specimens. We have identified intrageneric variations in the antennae shape, type of setation, coloration and even in the male genitalia of several other Leptolycini genera, which supports our statement above that the variation in the antennae and the pronotal mammiform tubercles in Baholycus are simply part of the intrageneric variation within Leptolycus. In Cessator, for example, whilst most specimens from the Puerto Rican bank have scaliform setae in the antennae, C. crypticus lacks this feature (Fig. 6D), and some of the specimens have the male genitalia with a phallobase several times longer than the median lobe (Fig. 7A,C) (C. obrienorum and C. luquillonis) whilst others present the median lobe and the phallobase similar in shape and length (Fig. 7D) (C. crypticus). The herein described Dracolycus have one of the species with the presence of mammiform tubercles in the pronotum (D. chupacabra), whilst the other species does not have these structures (D. marshalli). In the Leptolycus of the Puerto Rican bank, there is a wide variation in color patterns of pronota, thorax and antennae shapes, in both adult males and their immature forms (see detailed discussion under L. heterocornis), further exemplifying the plasticity of several features within genus level. Even leaving the taxa originally described in Baholycus from Hispaniola out of our examples, there is great variation in the antennae shape just in the Puerto Rican bank Leptolycus: L. heterocornis possesses a filiform antennae, with the inner margins crenulate (Fig. 13E–H), whilst L. viensis, and L. flavicollis have their antennae parallel-sided with no or only weak crenulation (Fig. 13A,B); L. falsoheterocornis has a tubular antenna and L. puellus has a subserrate antennae (Fig. 13C,D, respectively); all mentioned species have bristle-like setation on their antennae, with the exception of L. puellus, which possesses a short decumbent setation throughout. On the other hand, all Leptolycus and Baholycus possess similar genitalia (see differential diagnosis section above in Leptolycus). Therefore, to allow the existence of Baholycus as a separate subgenus in the Leptolycus we would have to describe a new subgenus for each of the variations found in the shape of the antennae and pronotum in all many Leptolycini, even though those variations are accepted to be within the expected range of intrageneric variation for all other mentioned taxa. Therefore, we synonymize the subgenus BaholycusBocak, 2001,new junior synonym with LeptolycusLeng and Mutchler, 1922. Leptolycus heterocornisLeng and Mutchler, 1922 Figs. 4, 7L–N, 13L–N, 14E–H, 15A–C, 16A–C, 17A–C, 20A Leptolycus heterocornisLeng and Mutchler, 1922: 430 fig.12 (in part, male of Leng and Mutchler); Kleine 1933: 18; Blackwelder 1945: 343. Miskimen and Bond 1970: 83. Leptolycus heterocornis var. flavicollisLeng and Mutchler, 1922:430 (in part); Blackwelder 1945: 343; Kazantsev 2009: 88. Leptolycus (Leptolycus) heterocornis;Kazantsev 2009: 89, 91 figs. 1, 4-17. Examined Type Material(n = 2). Holotype: 1: Aibonito, Porto Rico/ July 14-17, 1914/H. G. Barber/ Holotype/ Am. Mus. Nat. Hist./ Dept. Zool./ No. 24525 (AMNH) [Photograph]. Paratype: 1: Caveym P. R.; May30, 31, 1915/ PARATYPE [in red]/ Am. Mus. Nat. Hist.; Dept. Zool.; No. 24528/ Paratype; 61029; USNM/ Leptolycus; heterocornis; Leng & Mutchler; Paratype (NMNH). Material Examined in Addition to Type Specimens. Adults (n = 34). Color morph 1: 3: PUERTO RICO; GuilarteFor. Res.; Hwy.131&158, July; 23, 1979LB.O’Brien (WIBF). 4: PUERTO RICO; GuilarteFor. Res.; Hwy. 131&158, July; 23, 1979GBMarshall (WIBF). 1: PUERTO RICO, Hwy.; 120, K13H8, Maricao; For. Res., July 26; 1979 G.B. Marshall (WIBF). 1: PUERTO RICO, Carib.; N.F., ElToroNegroD.; K19H9, Hwy. 143, July; 22. 1979GB. Marshall (WIBF). 2: ToroNegro; PuertoRico; VII.1977 (NMNH). 1: PUERTO RICO: El Verde St.; 18.3218ºN, 65.8270ºW; 05JULY2017, 1,225 ft; Ferreira & Spiessberger; near FIT#5, sweeping/ WIBF 051532 (WIBF). 1: Insular Forest; Maricao, P.R.; III-29-1947/ J. A. Ramos; Collector (WIBF). 1: Adjuntas, P.R.; June 8-13, 1915 (AMNH). 1: PUERTO; Rico 8mi; E Mayaguez; II-9-1969/ Collectors: L&; C.W.O’Brien (WIBF). 1: Puerto Rico, Carib.; N.F., ElToroNegroD.,; Hwy.143, K18H6, 7-;22-1979CW. O’Brien (WIBF). 1: PUERTO RICO; GuajatacaFor.Res.; July 27, 1979; C.W.O’Brien (WIBF). 1: PUERTO RICO. Hwy.; 120. K13H8, Maricao; For.Res., July25,; 1979 L.B. O’Brien/ WIBF 051541 (WIBF). 1: Maricao Forest; 2-3,000 ft. P.R.; May 30-June 2, ‘38; Darlington (MCZ). 1: Acc # 203-47; Toro Negro, P.R.; Nov. 14, 1947 (UNIVERSITY OF PR). 1: slip says; Sandos [illegible]; #4845; vial label; 13 II 669/ Villalba PR; 26. Oct. 33; RGOakley/ Unknown Host; I.G. Finca; Villalba, P.R.; Cp;/ 26 Oct. 33; R.G. Oakley; 669 (NHNM). Color Morph 2: 4: PUERTO RICO: Rio Grande Mun.; El Verde Research Station; 18º19.376’N 65º48.997’W; 17JuL2007 sweeping; coll. M. Gimmel [one specimen WIBF 051540] (WIBF). 1: PUERTO RICO: Rio Grande Mun.; El Verde Research Station; 18º19.376’N 65º48.997’W; 16JuL2007 sweeping; coll. M. Gimmel (WIBF). 1: PUERTO RICO: El Verde Sta; 18.3218ºN, 65.8270ºW; 1-4JULY-2017, 1,225 ft; Ferreira & Spiessberger; near FIT 5, sweeping (WIBF). 1: PUERTO RICO: El Verde St.; 18.3218ºN, 65.8270ºW; 05JULY2017, 1,225 ft; Ferreira & Spiessberger; near FIT#5, sweeping/ MTEC 023636 (WIBF). 1: PUERTO RICO: El Verde Sta; 18.3218ºN, 65.8270ºW; 19JUNE-08JULY2017, 1,225 ft; E.L. Spiessberger &; V.S. Ferreira & Spiessberger Malaise#5 (WIBF). Color Morph 3: 4: El Yunque; c. 3,000 ft, P.R., May 1938; Darlington (MCZ). 1: El Yunque Sta.,; Luquillo Forest,; P.R. VII 10-16, 1969; H. & A. Howden/ WIBF 051539 (CNC). 1: El Yunque P.R.; Mar. 20-22, 1954/ J. Maldonado; S. Medina Colls. (AMNH). 1: El Yunque P.R.; Feb. 17-1925/ H.L. Dozier; Collector (NHMN). Color Morph 4: Yellow pronotum: 1: PUERTO RICO. Carib.; N.F., Mt.Britton.; Trail, July19 1979; G.B. Marshall/ WIBF 051538 (WIBF). Larvae (n = 20). Color morph 1: 1: PUERTO RICO: El Verde Field Station; July 2018; T. Sanchez & K. Noa Reyes (WIBF). 3: Puerto Rico: Aguas Buenas; Forest at Aguas Buenas; cave 250m, v: 7-17: 1973; leg S. Peck – Ber #; 265. FM (HD) #73-4, Ber:; 93 Kg., 171 l (WIBF). 12: PUERTO RICO; El Verde Research Station; ridge tops in forest; 15 MAY – 04 NOV 1996; E. Nazaro colr.; pitfall traps (WIBF). 1: PUERTO RICO: El Verde Field Station; July 2018; T. Sanchez & K. Noa Reyes/ WIBF 041693 (WIBF). Color morph 2: 1: PUERTO RICO: Naguabo; El Yunque Nat. Forest, S part; La Sabana recr. area 5.6 km N of Río Blanco at rd.PR191; 18°16.1’N, 65°47.6’W, 510 m, 21.vi.-2.vii.2016; Deler,Fikáček & Seidel lgt. PR03 [WIBF 051506] (WIBF). Color morph 3: 1: PUERTO RICO: Naguabo; El Yunque Nat. Forest, S part; 5.4 km N of Río Blanco, rd.PR191; at the gate,18°15.9’N,65°48.1’W; 510 m, 20–24.vi.2016; Deler, Fikáček & Seidel lgt. PR01 [WIBF 051507] (WIBF). 1: PUERTO RICO: Río Grande; El Yunque Nat. Forest, N part; 9.1 km SSE of Río Grande, tk. from Mt. Britton; to Mt. El Yunque 18°18.4’N, 65°47.53’W; 925 m, 22.vi.2016; Deler, Fikáček & Seidel PR06 [WIBF 051537] (WIBF). Diagnosis of Adult Males. Leptolycus heterocornis is one of the most distinctive Leptolycini species in Puerto Rico. Leptolycus heterocornis has a wide color variation in both adult males and immature forms (see color variability discussion after diagnosis), but the species can be easily separated from all other Leptolycus from the Puerto Rican bank by the following characters: the pronotum distinctly constricted medially, with lateral margins indistinct and only weakly carinated, seemingly absent (Fig. 13E–H) (vs. pronotum not distinctly constricted in other Leptolycus species, with lateral margins ranging from strongly to weakly developed in other species of the genus), the absence of a pronotal median posterior fovea (Fig. 13E–H) (vs. shallow median pronotal fovea present in L. puellus, L. viensis and L. flavicollis), the very long antennae and legs, with an overall slender distinct body type (Fig. 13E–H) (vs. variable in other species, if slender not as distinctive as in L. heterocornis and not possessing the same combination of characters as mentioned here), the antennae dorsoventrally strongly compressed and filiform (Fig. 13E–H) (vs. tubular in L. falsoheterocornis, subserrate in L. puellus), the inner margins of the antennae irregularly crenulate, sometimes strongly crenulate (Fig. 13E–H) (vs. weakly crenulate in L. flavicollis and L. viensis; entire in L. puellus and L. falsoheterocornis), and the interconnecting membranes in the apex of antennomere IV to the base of XI yellow, translucid, distinctly visible (Fig. 13E–H) (vs. membranes not conspicuous in L. viensis and L. puellus). Length (pronotum + elytra): 2.68–3.80 mm. Width (across humeri): 0.78–0.93 mm. Diagnosis of Larvae. Leptolycus heterocornis larvae can be easily separated from the other species of Leptolycus from the Puerto Rican bank by the subquadrate shape of its thoracic segments (Fig. 16A–C) (vs. transverse, subtrapezoidal or anteriorly round in other species), the unique color pattern, with the metanotum yellow or dark brown and abdominal segment V dorsally predominantly yellow (Fig. 16A–C) (vs. different color combination in other species) and by the strongly developed mandibles (Fig. 17A–C) (vs. weakly or moderately developed in other species). Length: 1.85–3.18 mm. Distribution. Puerto Rico (Fig. 20A). Incorrectly recorded from St. Croix (see distribution section under the genus redescription). Intraspecific Variation in L. heterocornis. Color patterns, when used as a main diagnostic characteristic, can be misleading in the taxonomy of aposematic beetles, especially in the Lycidae. The case with L. heterocornis is an example of this, and very well represents the limitations of a color-based taxonomic system, which was widely used as the main diagnostic character for several Neotropical Lycidae in the past. After our thorough and exhaustive examination of the external and male genitalic morphology of several specimens possessing the diagnostic characteristics mentioned above, we did not find any characteristics, other than the color variation, to support the separation of these color morphs into different species. In addition to color patterns, minor variations were observed in the proportion of antennomeres, pronotum shape, angles of the gular sutures, and mouth-cavity region, and in the shapes of the tips and proportions of the median lobe of the male genitalia. However, these variations were found in a nonconsistent manner across the different color variants, and they seemed more related to the ways that the specimens were preserved (pinned or pointed) and how old they were rather than constant and consistent morphological differences. Our findings and conclusions that color variations should not be used for defining species concepts were further corroborated by our DNA barcoding methods, in which larvae morphotypes 1–3, (Fig. 15A–C) matched together with maximum support level to our adults of Leptolycus heterocornis (Fig. 4). In the adult males of L. heterocornis, color varied in the following structures: prothorax being completely brown (Fig. 13E,F,H) or golden-yellow (Fig. 13G); the scutellar shield being yellow (Fig. 13F), golden-yellow (Fig. 13G), or brown (Fig. 13E,H), the meso- and metathorax brown (Fig. 13E,H), yellow (Fig. 13F), or golden-yellow (Fig. 13G). In the larvae, the variation ranged from the thorax and abdomen completely dark brown, with abdominal segment V mostly yellow, only medially dark brown (Fig. 15A–C), to a color pattern of metathorax and abdominal segment yellow, with the remaining of the thorax and abdomen brown or dark brown (Fig. 15A,B). Since color has been traditionally used for separating species in the Lycidae, we opted to present the examined material section discriminating these color variants to illustrate the group’s diversity and to avoid further misunderstanding in the taxonomy of the group in this species complex. Nomenclatural and Taxonomic Notes on the Identity of L. heterocornis and other Puerto Rican Leptolycus Species. At the time of Leptolycus heterocornis’ original description by Leng and Mutchler (1922), the authors inadvertently had in their type series three distinct species, which are nowadays referred to as Leptolycus heterocornis, Leptolycus puellus and Leptolycus albicauda. The specimen determined as the holotype of Leptolycus heterocornis by Leng and Mutchler (1922: 431: ‘Holotype male, No. 24525’), remains the sole name-bearer of that species (ICZN 1999, Art. 73.1). Following that, in our study, we re-diagnosed and validated that species concept based on the study of the morphology of the photograph of the holotype at the AMNH and several other specimens we had available at hand (see differential diagnosis section under this species). The specimen that was identified by Leng and Mutchler as the allotype of L. heterocornis (Leng and Mutchler 1922: 431: ‘...allotype female, No. 24526’) was examined by Miller (1991: 242) in his PhD dissertation, on which he compared the allotype No. 24526 with other AMNH specimens, including the holotype of L. heterocornis. At the time of his dissertation, Miller concluded that some of the specimens that have been historically included in the L. heterocornis type series should be rendered a separate species from L. heterocornis, due to variations in the antennae shape and pronotum (see differential diagnosis under L. puellus). Unfortunately, Miller’s dissertation was never published, and despite the existence of specimens identified with the name that he proposed for that species in certain scientific collections (Leptolycus effeminatus Miller in litteris), his proposed name remains invalid. Kazantsev (2009), in his publication on the Leptolycini of Puerto Rico, described a new Leptolycus species—Leptolycus puellus—that fits the species concept proposed by Miller in his dissertation (Leptolycus effeminatus). Kazantsev donated one of the paratypes of L. puellus to VSF, and the authors were able to compare the specimen with Leng and Mutchler’s L. heterocornis paratypes Nos. 24527 and 24528, which have been originally compared with Leng and Mutchler’s allotype of L. heterocornis (specimen No. 24526 in the AMNH). Even though we never examined the allotype of L. heterocornis, based on these cross-comparisons of specimens, we concluded that the allotype fits the characteristics of L. puellus, the species named by Kazantsev in 2009, which is the valid name for the species. Still in the original description of L. heterocornis, Leng and Mutchler mentioned the existence of a variety in their examined series, which they named Leptolycus heterocornis var. flavicollis. They described the variety as follows: ‘This variety agrees with the typical form [of Leptolycus heterocornis], except that the color of the thorax and scutellum is yellow. Holotype female, No. 24529.’ (1922: 431). Just like the above case with L. puellus, Miller (1991) studied the holotype of L. heterocornis var. flavicollis altogether with several other specimens and concluded that the variety should be treated as a separate species from L. heterocornis, proposing to elevate the variety to the level of species, which was never concretized because his dissertation remains unpublished. In Kazantsev’s (2009) publication, to the moment the most comprehensive study on the Leptolycini of Puerto Rico, under the synonimical table of L. heterocornis, Kazantsev mentioned the examination of the holotypes of L. heterocornis (specimen No. 24525) and L. heterocornis var. flavicollis (specimen No. 24529), and concluded that the latter was simply a color variant of L. heterocornis, rendering it the junior synonym of the latter, as follows: (page 88) ‘A specimen with yellow ochre pronotum was designated Holotype of L. heterocornis var. flavicollis Leng et Mutchler, 1922, which is hereby put in synonymy of L. heterocornis Leng et Mutchler, 1922’. Immediately after the taxonomic treatment of L. heterocornis in Kazantsev’s 2009 paper, the author described and named the new species Leptolycus albicauda. The description of that species was only based on a singleton deposited in Kazantsev’s personal collection, and no photos of the habitus of the species have been provided in formal literature. Even though no photographs of L. albicauda were available, the description and diagnosis of the species are very distinctive and precise, and a comparison of these with the photograph of the damaged holotype of L. heterocornis var. flavicollis in the AMNH on-line database allowed the immediate association of L. albicauda with L. heterocornis var. flavicollis by us. Miller also had the opportunity to study L. heterocornis var. flavicollis specimens (including the holotype, specimen No. 24529), having discussed his findings in his dissertation. Of the three specimens of L. heterocornis var. flavicollis studied by Miller (1991), we had firsthand access to two of them and access to the photograph of the holotype on the AMNH website (specimen No. 24529). After comparing our findings with Miller’s notes, we agreed with Miller’s opinion that L. heterocornis var. flavicollis should be treated as a separate species from L. heterocornis (see differential diagnostic under L. flavicollis), and we were inclined to think that L. albicauda was indeed a synonym of L. flavicollis. However, since the type specimen of L. albicauda has never been studied by any researchers other than its original author, we were unsure of our conclusions. To mitigate our doubts regarding the true identities of L. flavicollis and L. albicauda and to finally determine which was the oldest name available for that species, VSF contacted S. Kazantsev and provided him dorsal and ventral photographs of one of the specimens in-hand which have been identified as L. albicauda and cross-identified with L. heterocornis var. flavicollis holotype on the AMNH website. Upon examination of the shared photographs, Sergey Kazantsev (pers. communication) confirmed that the specimen that we imaged and sent to him matched with his holotype of L. albicauda. Therefore, based on this series of events and multiple direct and indirect comparisons and confirmations, we can confidently conclude that Leptolycus albicaudaKazantsev, 2009 is indeed a new junior synonym of Leptolycus var. flavicollisLeng and Mutchler, 1922, which is herein elevated to Leptolycus flavicollisLeng and Mutchler, 1922new status (see differential diagnosis under this species). Leptolycus flavicollisLeng and Mutchler, 1922 New Status Figs. 7H, 13A, 14A, 19A Leptolycus heterocornis var. flavicollisLeng and Mutchler, 1922: 430 fig.12 (in part, female of Leng and Mutchler); Kleine 1933: 18; Blackwelder 1945: 343. Leptolycus (Leptolycus) heterocornis;Kazantsev 2009: 89, 91 figs. 1, 4–17. Leptolycus (Leptolycus) albicaudaKazantsev, 2009: 88, 91 figs. 18, 19. New junior synonym. Examined Type Material. Holotype: 1: PUERTO RICO: Aibonito, July 1914, H. G. Barber (AMNH) [Photograph]. Material Examined in Addition to Type Specimen. Adults (n = 4). 1: PUERTO RICO; GuilarteFor.Res.; Hwy.131&158, July; 23, 1979LB. O’Brien/ WIBF051543 (WIBF). 1: Consumo P.R.; 12-Oct-1937; J.A. Ramos; Collector/ WIBF051542 (EPRL). 1: PUERTO RICO: Rio; Grande, El Yunque, Mt.; Britton summit. 941m; 4-12.VIII.1999, P.W.; Kovarik, collector FIT (WIBF). 1: PUERTO RICO: Rio Grande Mun.; El Verde Research Station; 18º19.376’N 65º48.997’W; 17Jul2007 sweeping; coll M. Gimmel (MLGC). Diagnosis of Adult Males. Leptolycus flavicollis can be easily separated from all other Leptolycus in the Puerto Rican bank by its unique color pattern, with its prothorax, mesothorax and humeral region of elytra yellow (Fig. 13A) (vs. variable in other species, but never with the present combination), the terminal segments of the abdomen (ventrite VIII and tergites IX and X) white (vs. terminal segments of the abdomen of other Leptolycus brown or black), the pronotum with a posterior median fovea weakly developed (Fig. 13A) (vs. fovea absent in L. heterocornis), the scutellar shield notched apically, with angles distinctly acute (Fig. 13A) (vs. scutellar shield entire in L. heterocornis, L. puellus), and the male genitalia with the median lobe 0.5× longer than the phallobase, which is oval (Fig. 7H) (vs. variable in other Leptolycus). Length (pronotum + elytra): 3.02–3.30 mm. Width (across humeri): 0.76 mm. Distribution. Puerto Rico (Fig. 19A). Immature and Females. Unknown. Taxonomic Note. See ‘Nomenclatural and taxonomic notes on the identity of L. heterocornis and other Puerto Rican Leptolycus species’ under L. heterocornis. Leptolycus puellusKazantsev, 2009 Figs. 4, 7K, 13D, 14D, 19A Leptolycus heterocornisLeng and Mutchler, 1922: 430 fig.12 (in part, male of Leng and Mutchler); Kleine 1933: 18; Blackwelder 1945: 343. Leptolycus (Leptolycus) heterocornis;Kazantsev 2009: 89, 91 figs. 1, 4–17. Leptolycus (Baholycus) puellusKazantsev, 2009: 91 figs. 20–23. Examined Type Material. Leptolycus puellus: Paratype: 1: PUERTO RICO: Sierra; de Luquillo 600 m; 3 5.IV.2005; S. Kazantsev leg./ [in red] Paratype; Kazantsev 2009/ WIBF 051546 (WIBF). Leptolycus heterocornis: Paratypes: 1: Aibonito, P.R.; July 14–17, ‘14/ [in red] PARATYPE/ Am. Mus. Nat. Hist.; Dept. Invert. Zool.; No. 24528/ WIBF 051547 (AMNH). 1: Aibonito, P.R.; July 14-17, ‘14/ [in red] PARATYPE/ Am. Mus. Nat. Hist.; Dept. Invert. Zool.; No. 24527/ [handwriting] Leptolycus; heterocornis; type L&M/ WIBF 051548 (AMNH). Material Examined in Addition to Type Specimens. Adults (n = 20). 11: PUERTO RICO; El Verde; statopm road; 2.ii.1989; S.A. Marshall (DEBU). 1: Maricao P.R.; 2-8 1936; Coll: J.A. Ramos (EPRL). 1: PUERTO RICO,; GuajatacaFor. Res.; July 27, 1979; C.W. O’Brien (WIBF). 2: PUERTO RICO; GuilarteFor. Res.; Hwy. 131&158, July; 23, 1979GBMarshall [one specimen WIBF 051549] (WIBF). 1: PUERTO RICO; CambalacheForest; Res., July28, 1979; C.W. O’Brien (WIBF). 1: PUERTO RICO: El Verde Field Station; sweeping; V.S. Ferreira colr.; 1-4 July/ WIBF 041665 (WIBF). 1: PUERTO RICO: El Verde Sta.; 18.3218ºN,65.8270ºW; 30JUNE2017, 1,225 ft; Ferreira & Spiessberger; near FIT 5, sweeping/ WIBF 051545 (WIBF). 5: PUERTO RICO: El Verde Sta; 18.3218ºN,65.8270ºW; 1-4JULY-2017, 1225 ft; Ferreira & Spiessberger; near FIT 5, sweeping [one specimen WIBF 051544] (WIBF). 3: PUERTO RICO: El Verde Sta; 18.3218ºN,65.8270ºW; 05JULY2017, 1225 ft; Ferreira & Spiessberger; near FIT#5, sweeping [WIBF 051534, WIBF051533, MTEC023635] (WIBF). 3: PUERTO RICO: Rio Grande Mun.; El Verde Research Station; 18º19.376’N 65º48.997’W; 17JuL2007 sweeping; coll. M. Gimmel (WIBF). 1: PUERTO RICO: Rio Grande Mun.; El Verde Research Station; 18º19.376’N 65º48.997’W; 16JuL2007 sweeping; coll. M. Gimmel (MLGC). Diagnosis of Adult Males. Leptolycus puellus can be separated from all Leptolycus from Puerto Rico by the following combination of characters: pronotum with lateral margins distinct, not constricted medially (Fig. 13D) (vs. lateral margins indistinct and strongly constricted medially in L. heterocornis), the presence of a shallow posterior median fovea in the pronotum (Fig. 13D) (vs. absent in L. heterocornis), the subserrate antennae, with the margins entire (Fig. 13D) (vs. tubular in L. falsoheterocornis and filiform in L. heterocornis and L. flavicollis, weakly crenulate in the first and moderately to strongly in the last two), the antennae in resting position surpassing the elytral apex (vs. not surpassing elytral apex in L. falsoheterocornis), the scutellar shield entire (Fig. 13D) (vs. scutellar shield apically notched in L. flavicollis and L. falsoheterocornis) and the male genitalia with the median lobe subequal in length with the phallobase (Fig. 7K) (vs. variable in other Leptolycus). Length (pronotum + elytra): 2.24–2.78 mm. Width (across humeri): 0.54–0.61 mm. Distribution. Puerto Rico (Fig. 19A). Immature and Females. Unknown. Note on the Intraspecific Variation of L. puellus. We observed that most specimens that have been collected in the past 10 y window (2011–2021) presented a more vibrant coloration, with their pronota, head, antennae and scutellar shield black instead of dark brown (as observed in the paratypes and in older specimens) and a predominance of elongate scaliform setae on their antennae (which was seemingly absent in older specimens). Some specimens presented variation in the coloration of the gular region (black or dark brown or yellow) and coloration of coxae (entirely black or partially yellow). These variations were present in specimens of different locations and collected in different moments and times by different collectors. After study of their external morphology, male genitalia and their COI sequences, the latter with 99% similarity or more in the sampled specimens (see Supp. Materials S3–S6 [online only]) (specimens WIBF 041665 and MTEC 023635 have brown antennae whilst specimens WIBF 05133 and WIBF051534 have black antennae, being one of each with their coxae with different colors) we are simply considering these to be in the spectrum of intraspecific variability for this species. Taxonomic Note. See ‘Nomenclatural and taxonomic notes on the identity of L. heterocornis and other Puerto Rican Leptolycus species’ under L. heterocornis. Leptolycus falsoheterocornis Ferreira and Ivie New Species Figs. 7J, 13C, 14C, 19A Zoobank LSID: urn:lsid:zoobank.org:act:15B29C22-4D1E-45B0- 93E2-E9810B515EF4 Type Material. Adults (n = 1): Holotype: 1: PUERTO RICO; GuilarteFor.Res.; Hwy.131&158, July; 23, 1979LB. O’Brien (WIBF). Etymology. The species has been given this name due to its superficial resemblance with L. heterocornis. The word falso comes from Latin and means false, deceitful. Diagnosis of Adult Males. Leptolycus falsoheterocornis can be separated from its congenerics from the Puerto Rican bank by the pronotum with lateral margins distinct, not constricted medially, and the presence of a shallow posterior median fovea in the pronotum (Fig. 13C) (vs. lateral margins indistinct and strongly constricted medially in L. heterocornis, fovea absent), the tubular antennae, not dorsoventrally compressed (Fig. 13C) (vs. variable in other Leptolycus, but always dorsoventrally flattened; filiform in L. heterocornis, L. viensis and L. flavicollis, subserrate in L. puellus), and the scutellar shield notched apically (Fig. 13C) (vs. notch absent in L. heterocornis). Description of Male. General coloration: Head, pronotum, scutellar shield, antennae, legs and abdomen brown (Fig. 13C); antennomeres XI and metatrochanters yellow; elytra bicolored, basal fourth yellow, remaining brown (Fig. 13C). Head: frons weakly developed, not bulgy (Fig. 13C). Maxillary palpomeres IV acuminate. Gular sutures basally connected by a bridge (Fig. 14C). Antennae tubular, not dorsoventrally compressed, bearing bristle-like erect setae, in resting position reaching elytral apical fourth (Fig. 13C); antennomere IV elongate, 3x the length of the sum of scape + pedicel (Fig. 13C). Pronotum transverse, not medially constricted, margins distinct, apical and basal margins developed; bearing weakly developed posterior median fovea (Fig. 13C). Scutellar shield posteriorly medially notched (Fig. 13C). Metatrochanters tubular, elongate, Median lobe of the male genitalia acuminate, 0.5× longer than phallobase, which is oval and posteriorly round (Fig. 7J). Length (pronotum + elytra): 2.86 mm. Width (across humeri): 0.64 mm. Distribution. The only known specimen is from Puerto Rico, from the Bosque Estatal de Monte Guilarte (Fig. 19A). Immature and Females. Unknown. Leptolycus viensis Ferreira and Ivie new species Figs. 7I, 13B, 14B, 15G, 16G, 17G, 18, 20B Zoobank LSID: urn:lsid:zoobank.org:act:5044EA7A-30A9-40A0- AB14-45A5B3285657 Examined Type Material. Adults (n = 41): Holotype: 1: BR. VIRGIN IS: Guana; 02 NOV 1990, 0-80 m; S. E. Miller colr.; at u.v. light [WIBF 051560] (BPBM). Paratypes: 3: BR: VIRGIN IS: Guana Is; Quail Dove Ghut; 13NOV-15DEC1992, 400 ft; Lio Wei Peng colr.; flight inter. trap #5 [WIBF 000720-722] (WIBF). 6: BR: VIRGIN IS: Guana Is; Quail Dove Ghut; 13NOV-15DEC1992, 600 ft; Lio Wei Peng colr.; flight inter. trap #5 [WIBF 000955-000960] (WIBF). 1: BR: VIRGIN IS: Guana Is; Quail Dove Ghut; 25DEC–25JAN1993, 600 ft; Lio Wei Peng colr.; flight inter. trap #13 [WIBF 000980] (WIBF). 1: BR: VIRGIN IS: Guana Is; Quail Dove Ghut; 24JULY–09 OCT 1994; M.A. & L.L. Ivie; flight intercept #5 [WIBF 021888] (WIBF). 1: BR: VIRGIN IS: Guana Is; Quail Dove Ghut; 12JULY-09 OCT 1994; flight intercept #13; M.A. & L.L. Ivie colr. [WIBF 021595] (WIBF). 2: BR: VIRGIN IS: GuanaIs; Quail Dove Ghut; 24-31OCT1992, 400ft.; M.A. & L.L. Ivie colr.; flight inter. trap #5 (WIBF 000680, WIBF 000681) (WIBF). 1: BRITISH VIRGIN IS:; Guana I; 13.vii.1988/ S.E. Miller & C.; O’Connell cols. (WIBF 013271) (WIBF). 2: BRITISH VIRGIN; ISLANDS; Guana Island/ 26-X-1997/ C.R. Bartlett; beating/sweeping (WIBF). 3: BRITISH VIRGIN IS:; Guana I: x.1989; V.O. Becker, Coll.; BISHOP Museum [WIBF 013272](BISHOP MUSEUM). 2: BRITISH VIRGIN IS:; Guana ‘beetletrap’; (flight intercept)/ July-Oct.1993; C. Bartlett &; J. Cryan [WIBF 018244, WIBF 018245] (NCSU). 1: BR. VIRGIN IS: Guana; 02 NOV 1990, 0–80 m; S. E. Miller colr.; at u.v. light (BPBM). 1: BR. VIRGIN IS: Tortola; Mt. SageNat. Pk. N. side; Mt.Sage, 1,550 ft, steps; 25 JULY-10 OCT 1994; M.A. Ivie & S.A. Bucklin [WIBF 023147](WIBF). 1: BR. VIRGIN IS: Tortola; Mt. SageNat. Pk. N. side; Mt.Sage, 1,550 ft, steps; 25 JULY-10 OCT 1994; flt. int. tr.#3 VIBFP [WIBF 022045] (WIBF). 3: VIRGIN IS: St. John; Est. Caneel Bay, Caneell; Hill. 240ft. 17DEC1992; 02JAN1993, VIBFP colrs; flight inter. trap #6 [WIBF 003780-003782, WIBF 004227] (WIBF). 1: VIRGIN IS: St. John; Est. Caneel Bay, Caneell; Hill. 240 ft. 02-14NOV; 1992. M.A. Ivie colr.; flight inter. trap #6 [WIBF 004173] (WIBF). 1: VIRGIN IS: St. Thomas; Est. Botany Bay; 29 JULY-15 OCT 1994; M.A. & L.L. Ivie [WIBF 022609] (WIBF). 1: VIRGIN IS: St. John; Est. Lameshur Bay; Reef Bay Trail, 5 ft; 27JUL-14OCT1994; M.A.& L.L. Ivie colrs. [WIBF 022735] (WIBF). 1: VIRGIN IS: St. John; Est. Lameshur Bay; Reef Bay Trail, 5 ft; 03JAN1993-06JULY1994; flight intercept #11 [WIBF 021998] (WIBF). 6: BRIT VIRGIN ISLS; Guana Island; July-October 1993; ‘Beetle-trap’/ Collected by; C. Bartlett &; J. Cryan [WIBF 017741, WIBF 017974, WIBF 017971, WIBF 017768, WIBF 017772, WIBF 017769] (NCSU). 1: VIRGIN IS: St. John; Est. Carolina, 250 ft; N.W. of Coral Bay; 18 MAY 1984; W.B. Muchmore colr.; ex ground litter [WIBF 051520] (WIBF). 1: on grass; St. John, V.I.; V. 29’40; D. Deleon/ Hopk. US; 33162.9 (NHNM). Females (n = 2): 1: VIRGIN IS: St. John; Est. Carolina, 250 ft; N.W. of Coral Bay; 18 MAY 1984; W.B. Muchmore colr.; ex ground litter [WIBF 051520] (WIBF). 1: VIRGIN IS: St. John; Lameshor Bay UPERS; 07MAY1984; W.B. Muchmore; base of roadside kapok (WIBF). Larvae (n = 24): 1: VI: St. John; Caneel Bay Est.; flight intercept #6, 73m; 17DEC-02JAN1993; VIBFP colrs. (WIBF). 2: VIRGIN IS: St. John; Francis Bay pond; 29Feb1989; W.B.Muchmore; base of lg. tree; on shore of pond (WIBF). 2: VIRGIN ISLANDS: St. John; Brown Bay, 11 March 1984; W.B. Muchmore colr.; litter and Calabash tree (WIBF). 1: VIRGIN IS: St. John; Hermitage; 03 MAY 1984; under donkey dung; W.B. Muchmore (WIBF). 1: VIRGIN IS: St. John; Catherineberg; 14MAY1984; litter between rocks; W.B. Muchmore (WIBF). 1: VIRGIN IS: St. John; Mahe Bay; 12MAR1984; W.B. Muchmore; under lg Seagrape; along road (WIBF). 2: VIRGIN IS: St. John; Bordeaux Mt.; 15MAY1984; ground litter among; trees & boulders; W.B. Muchmore (WIBF). 1: VIRGIN IS: St. John; Lameshur Bay UPERS; 07MAY1984; W.B. Muchmore; base of roadside kapok (WIBF). 1: VIRGIN IS: St. John; Kingshill, south; 08MAY1984; W.B. Muchmore; ground litter (WIBF). 6: VIRGIN IS: St. John; Est. Hermitage,; ruins; 20 JUN 1980/ base of giant; dildo cactus; Cephalocereus (WIBF). 1: VIRGIN IS: St. John; Trunk Bay; 09 JUN 1980; W.B. Muchmore/ at base of tree; and anthirium (WIBF). 2: VIRGIN IS: St. John; Lameshur Bay; Gray Gut; 12 JAN 1980; base of large; tamarind (WIBF). 1: VIRGIN IS: St. John; Cinnamon Bay Nature Trail; 06 JUN 1980; ex kapok buttreases; W.B. Muchmore (WIBF). 1: VIRGIN IS: St. John; Hawknest Bay; 28 FEB 1984, W.B. Muchmore; on and around huge boulder; in gut (WIBF). 1: VIRGIN IS: St. John; Windberg ruins; 08 OCT 1979; ex litter; W.B. Muchmore (WIBF). Etymology. This species has been given the name viensis after its distribution in the Virgin Islands (VI), where it is apparently endemic. Diagnosis of Adult Males. Leptolycus viensis is the only Leptolycus species in the Virgin Islands. This species can be immediately recognized among the other Leptolycus from the Puerto Rican bank by its transverse pronotum bearing a median posterior fovea (Fig. 13B) (vs. pronotum longer than wide in other Leptolycus species; median posterior fovea absent in L. heterocornis, L. flavicollis and L. puellus), the relatively short antennae, in rest position not surpassing the mid-elytral portion (Fig. 13B) (vs. long antennae, surpassing mid-elytra in all other species), the overall shortened antennomeres IV–XI (Fig. 13B) (vs. antennomeres IV–XI longer in other species), the metatrochanters shortened, semi triangular (vs. metatrochanters elongate and tubular in all other species) and the scutellar shield posteriorly medially notched (Fig. 13B) (vs. posterior margin of scutellar shield entire in L. heterocornis and L. puellus). Description of Male. General coloration: Head, pronotum, scutellar shield, antennae, legs, and abdomen brown (Figs. 13B, 14B); antennomeres XI yellow; elytra bicolored, basal third yellow, remaining brown (Fig. 14B). Head: frons weakly developed, not bulgy (Fig. 14B). Maxillary palpomere IV acuminate. Gular sutures basally connected by a bridge (Fig. 14B). Antennae filiform, dorsoventrally compressed, bearing bristle-like erect setae (Fig. 13B); in resting position not surpassing mid-elytral (Fig. 13B); antennomere IV elongate, ca. 2.5× the length of the sum of scape + pedicel (Fig. 13B). Pronotum transverse, not medially constricted, margins distinct, apical, and basal margins developed; bearing weakly developed posterior median fovea (Fig. 13B). Scutellar shield posteriorly medially notched (Fig. 13B). Metatrochanters subtriangular, short. Median lobe of the male genitalia acuminate, 1/3 longer than phallobase, which is shortened and posteriorly irregularly shaped, roundish (Fig. 7I). Length (pronotum + elytra): 2.78–3.62 mm. Width (across humeri): 0.73–0.85 mm. Diagnosis of Larvae. Leptolycus viensis larvae can separated from the other species of Leptolycus from the Puerto Rican bank by the subtrapezoidal shape of its pronotum, widest at the apex (Fig. 15G) (vs. subquadrate in L. heterocornis or transverse in other species), the nasale apically straight (Fig. 15G) (vs. bisinuate in L. heterocornis and other species), the weakly developed mandibles (Fig. 16G) (vs. strongly developed in L. heterocornis, variable in other species), the apparent absence of labial palps (Fig. 16G) (vs. labial palps minute, but clearly visible in L. heterocornis and other species) the unique color pattern, with the metanotum and abdominal segments IV and V predominantly yellow, but medially light brown and the abdominal segment IX yellow (vs. variable in other species). Diagnosis of Adult Female. The extreme paedomorphic adult female of Leptolycus viensis can be separated from the larva by its coloration uniform throughout, pale yellow (Fig. 18A) (vs. bicolored body in the larvae), the completely modified head, characterized by the nasale in dorsal view clearly bisinuate (Fig. 18D,E) (weakly bisinuate in the larva), the wide ventral plate of the head medially bearing a deep and wide V-shaped notch, with apices apically divergent (Fig. 18E) (vs. ventral plate narrowing medially towards apex, apically bearing a shallow v-shaped notch), the absence of maxillae and labial palpi (Fig. 18C,E) (vs. presence of maxillae in the larvae), the presence of only one pleurite (Fig. 18B) (vs. two pleurites in the larvae) and the absence of a pygopodium in abdominal tergite X (Fig. 18B,C) (vs. pygopodium present in the larvae). Distribution. This species is apparently endemic to the Virgin Islands, being found in St. John, St. Thomas, Guana and Tortola (Fig. 20B). Diagnosis of Unidentified Leptolycus Larvae Leptolycus sp. 1 Figs. 15D, 16D, 17D Examined Material (n = 7). 7: PUERTO RICO; El Verde Research Station; ridge tops in forest; 15 MAY-04 NOV 1996; E. Nazaro colr.; pitfall traps (WIBF). Differential Diagnosis. Leptolycus sp. 1 can be distinguished from other Leptolycus from Puerto Rico by their transverse thoracic segments (Fig. 15D) (vs. subquadrate in L. heterocornis and subtrapezoidal in L. viensis), the prominent antennal insertions (Fig. 15D) (vs. less conspicuous in other species) and by its creamy coloration (Figs. 15D, 16D, 17D) (vs. predominantly brown or dark brown in other species, with yellow patches throughout the body). Length: 2.41 mm. Leptolycus sp. 2 Figs. 15E, 16E, 17E Examined Material (n = 2). 1: PUERTO RICO: Patillas, Bosque Estatal Carite 9.6 km N of Patillas; Charco Azul tourist area; 18°5.5’N 66°1.9’W; 600 m; 23.vi.2016; Deler, Fikáček & Seidel PR09/ sifting of small accummulations; of leaf litter and thin layer of leaf; litter with lots of fungal mycelia; in a secondary forest on the slope above the dam [WIBF 051519] (WIBF). 1: PUERTO RICO: Arecibo; Bosque Estatal Río Abajo, 1.3 km SWW of the campsite at Cueva de las Aguas, 18°19.4’N; 66°43.0’W; 330 m, 27.vi.2016; Fikáček & Seidel lgt./sifting of leaf litter and mosses; on trunks and stones in sparse; lowland forest on limestone bedrock/ side pools of a stony stream with many decaying leaves (WIBF). Differential Diagnosis. Leptolycus sp. 2 is unique among the other Leptolycus larvae by having its body onisciform (Fig. 15E) (vs. parallel sided in other Leptolycus, with exception of Leptolycus sp. 3), the transverse thoracic segments (Figs. 15E, 16E) (vs. subquadrate in L. heterocornis and subpentagonal in L. viensis), the prominent antennal insertions, which are strongly developed in this species (Figs. 15E, 16E, 17E) (vs. variable in other species), and by the unicolored body, which is dark brown (Figs. 15E, 16E, 17E) (vs. bicolored in all other species, with the exception of Leptolycus sp. 1). Length: 2.72 mm. Leptolycus sp. 3 Figs. 15F, 16F, 17F Examined Material (n = 2). 2: PUERTO RICO: San Germán; Bosque Estatal Maricao, 8.6 km N of Sabana Grande; 18°8.0’N 66°59.5’W; 820 m 28.vi.2016; Fikáček & Seidel lgt. PR16 [WIBF 051503] (WIBF). Differential Diagnosis. Leptolycus sp. 3 is very similar to Leptolycus sp. 2, being readily separated from the former by its unique color pattern, with the side margins of the metathorax and the abdominal segment IV laterally yellow (Figs. 15F, 16F, 17F) (vs. specimen completely dark brown in Leptolycus sp. 2). Length: 4.29 mm. A Key to the Adult Male Leptolycus of the Puerto Rican Bank 1. Pronotum distinctly constricted medially, with lateral margins indistinct and only weakly carinated, never bearing a median posterior fovea. Overall slender individuals, with relative long legs, antennae, and elytra Leptolycus heterocornisLeng and Mutchler, 1922 complex 1.’ Pronotum with lateral margins distinct, not constricted medially, posteriorly bearing a median shallow fovea or fovea absent. Overall body type variable, ranging from slender to stout and short bodied individuals 2 2. Prothorax, mesothorax and humeral region yellow. Terminal segments of the abdomen entirely whitish. Scutellar shield notched apically, angles distinctly acute Leptolycus flavicollisLeng and Mutchler, 1922 2.’ Prothorax and mesothorax black, elytral apical third yellow. Terminal segments of the abdomen dark brown. Scutellar shield entire or shallowly notched apically, if notched, angles not acute 3 3. Antennae serrate bearing short decumbent setae; pronotum not bearing a median posterior fovea Leptolycus puellusKazantsev, 2009 3.’ Antennae filiform, bearing long bristle setae; pronotum bearing a median posterior fovea, sometimes weakly developed 4 4. Antennomeres IV–X tubular; antennomere IV nearly twice the length of the precedent combined; elytral costa I only distinctly visible in basal third of elytra; preantennal gibbosity pits weakly developed Leptolycus falsoheterocornis Ferreira and Ivie new species 4.’ Antennomeres IV–X flat and wide, dorsoventrally strongly flattened; antennomere IV 1/3 longer than the sum of the precedent combined; elytral costa I only distinctly visible up until middle third of elytra, fused apically with costa II; preantennal gibbosity pits strongly developed Leptolycus viensis Ferreira and Ivie new species TainopteronKazantsev, 2009 Figs. 5G, 6G, 7G, 19B TainopteronKazantsev, 2009: 88, 92. Differential Diagnosis of Adult Males. Tainopteron can be distinguished from all other Leptolycini by the following characters: the pedicel and antennomere III subequal in length (Fig. 5G, 6G) (vs. antennomere III multiple times longer than pedicel in Cessator), the dorsoventrally compressed antennae bearing bristle-like erect setae (vs. variable across the different genera, but tubular in Cessator and not bearing bristle-like setae), the strongly transverse pronotum (Fig. 5G) (vs. longer than wide in Dracolycus, Dominopteron and some Leptolycus), with bisinuate posterior angles and a strongly demarcated median fovea (Fig. 5G) (vs. posterior angles with variable combination in other Leptolycini), the scutellar shield subquadrate, apically notched (Fig. 5G) (vs. shape and presence of notch variable in other genera) and a trilobate male genitalia, with a sword-like median lobe, which is apically blunt, the co-adapted small and apically acuminate parameres and a phallobase with the posterior margin round (Fig. 7G) (vs. variable across the other genera). Type Species. Tainopteron milleriKazantsev, 2009 (by monotypy). Distribution. Tainopteron is only known to occur in Puerto Rico, in the El Yunque National Forest region (Fig. 19B). Biology. Except for the type series, all other known Tainopteron specimens were collected using Flying Intercept Traps, suggesting that members of this group are active individuals. Specimens from the type series were collected in low strata foliage and grass (Kazantsev 2009) in a very similar manner and kind of habitat as many other Leptolycini. Tainopteron milleriKazantsev, 2009 Figs. 5G, 6G, 7G, 19B TainopteronKazantsev, 2009: 88, 92. Examined Material (n = 12): 3: PUERTO RICO: El Verde Sta.; 18.3218ºN,65.8170ºW; 28–29JUNE2017, 1,225 ft; V.S. Ferreira &; E.L. Spiessberger, FIT6 (WIBF) [WIBF051550, WIBF051551]. 3: PUERTO RICO: El Verde Sta.; 18.3236ºN,65.8183ºW; 18JUNE–8JULY2017, 1,033 ft; E.L. Spiessberger &; V.S. Ferreira, Malaise #4 (WIBF). 1: PUERTO RICO: El Verde Sta.; 18.3218ºN,65.8170ºW;19JUNE–08JULY2017 1,225 ft; E.L. Spiessberger &; V.S. Ferreira, Malaise #5 (WIBF). 1: PUERTO RICO: El Verde Sta.; 18.3236ºN,65.8170ºW; 20–23JUNE2017 1,033 ft; V.S. Ferreira &; E.L. Spiessberger, FIT4/ WIBF051524 (WIBF). 1: PUERTO RICO: El Verde Sta.; 18.3230ºN,65.8183ºW; 23JUNE2017, 1,010 ft; V.S. Ferreira &; E.L. Spiessberger, FIT 3 (WIBF). 1: PUERTO RICO: El Verde Sta.; 18.3230ºN,65.8183ºW; 14JUNE–07JULY2017 117 ft; V.S. Ferreira &; E.L. Spiessberger, Malaise1 (WIBF). 1: PUERTO RICO: Rio; Grande, El Yunque, Mt; Britton summit, 541 m; 4–12AUG1999 P.W.; Kovarik, FIT (WIBF). 1: PUERTO RICO: Naguabo; El Yunque Nat. Forest, S Part; 4.9 Km N iof Rio Blanco,; 18º15.8’N, 65º47.3’W, 495 m; 24.vi.-2.vii.2016; Fikáček & Seidel lgt. PR11/ flight intercept trap at the margin; of the rainforest in an area with many flowering Etilingera elatior; plants (NMPC). Rediagnosis, Distribution and Biology of Adult Males. This is a monotypic genus, and the information is the same as given under the genus above. Length (pronotum + elytra): 1.45–1.75 mm. Width (across humeri): 0.46–0.58 mm. An Identification Key to the Puerto Rican Bank Leptolycini Adult Genera 1. Pedicel small, multiple times shorter than antennomere III CessatorKazantsev, 2009 1.’ Pedicel and antennomere III subequal in length 2 2. Elytra remarkably dehiscent, ligulate, tapered towards apex; metatrochanters distinctly elongate and tubular; long legs and slender body LeptolycusLeng and Mutchler, 1922 2.’ Elytra moderately or weakly dehiscent, not ligulate, similar throughout; metatrochanters short, semi triangular; legs and body type variable, but not as slender as above 3 3. Pronotum strongly transverse, laterally imarginated; elytral costae weakly developed, fading towards apex; antennae bearing long-bristle like setae; scutellar shield subquadrate, apically notched; short and small-sized specimens (<1.75 mm long) TainopteronKazantsev, 2009 3.’ Pronotum longer than wide, all margins distinct; elytral costae strongly developed and visible throughout, fused apically; antennae bearing short decumbent setae; scutellar shield cordiform; long and big-sized specimens (>4.88 mm long) Dracolycus Ferreira and Ivie new genus An Identification Key to the Known Puerto Rican Bank Leptolycini Larvae and Female Genera 1. Anterior portion of the head bearing a pair of facing forward projections; body unicolored 2 1.’ Anterior portion of the head not bearing projections; body unicolored or bicolored 3. Leptolycus 2. Anterolateral portions of the head strongly stalked (Figs. 8D–F); body longitudinal line absent (Figs. 8D–F); long erect setae absent Dracolycus 2.’ Anterolateral portions of the head weakly stalked (Figs. 8A–C); presence of a distinct and strongly visible body longitudinal line (Figs. 8A–C); presence of long erect setae arising from the tergites and pleurites Cessator 3. Abdomen only bearing one pleurite, ventrites entire, not lobed (Fig. 18B); maxillae and labial palps seemingly absent (Fig. 18C,E); absence of a pygopodium (Fig. 18A–C) Leptolycus female 3.’ Abdomen bearing two pleurites, ventrites laterally lobed (Fig. 16A–F); maxillae and labial palps present (Fig. 15A–F); pygopodium present (Fig. 17A–F) Leptolycus larva Discussion DNA barcoding is a powerful tool in biodiversity studies. The versatility and relative low cost of this approach allows its use in a variety of contexts, ranging from biodiversity inventories (e.g., Telfer et al. 2015), identification and delimitations of cryptic species (e.g., Etzler et al. 2013), phylogenetic analyses (e.g., Sikes and Mullen 2021) to life-stages associations (e.g., Levkanikova and Bocak 2009, Etzler et al. 2014, Zhai et al. 2017). While the limitations of the method have been the subject of various debates (see references below), this approach continues to be widely accepted and used by the academic community, especially because of its extensive documentation and relative ease of interpreting results. In Lycidae, the implementation of this method has been used in a variety of studies in the past two decades, with different levels of success. Some studies reported very little differences in DNA sequences of taxa that were morphologically distinct (Kusy et al. 2018b), mixed results (Bocek et al. 2019) or a high level of congruence between barcodes and morphology (Jiruskova and Bocak 2015, Li et al. 2018b). Despite some controversy on the use of DNA barcode in a phylogenetic context, this approach works well in providing the identification and association of immatures with their adult forms (see Lekvanikova and Bocak 2009, for example), allowing the development of a framework to quickly enhance the taxonomic knowledge on the immature forms of beetles of groups that are poorly known and rarely collected, as are most lycid groups in the New World. While life-stage association through comparison of DNA sequences via barcoding has become increasingly popular in the past 25 y, newer alternative methods are promising even faster, cheaper and more efficient results than Sanger-based barcoding (e.g., MinION) (Srivathsan et al. 2018, 2019), but such methods require the sample materials to be freshly collected or that it have been preserved in ethanol 95% and kept at low temperatures (–20°C or colder). Such requirements rule out most of the specimens in many of the scientific collections, especially historical material and immatures of beetles, the latter which are often boiled prior their final preservation in Ethanol 70% or in other preserving agents that have a degrading action on genetic material (see Costa et al. 1988, Reiss et al. 1995, Ferreira and Costa 2017). For understudied groups like the Leptolycini, the availability of material for traditional taxonomic and systematic research in scientific collections is already small, and the existence of DNA grade material even smaller. In our study, most of the available specimens came from legacy material deposited in scientific collections, and nearly all larvae came from a single institution. The DNA quality specimens of the immature forms of Leptolycini were the result of thousands of dollars and months of combined fieldwork in Puerto Rico from VSF and collaborators, and networking with other researchers from other institutions, yielding very few viable specimens for barcoding (we were only able to obtain sequences of 13 immature individuals). The development of this project and the implementation of barcoding methods was only possible to occur thanks to the availability of dozens of specimens that have been collected, sorted, labeled, tentatively identified as a Lycidae and finally permanently deposited in a scientific collection that we had access to. The availability of these specimens, in particular the L. viensis larvae and adults from the Virgin Islands—the only known Lycidae species from St. John, St. Thomas and Guana Island (Fig. 20B)—provided to us a direction as to what to look for in Puerto Rico, which resulted in the successful cross-matching and life-stage associations based on biogeography and DNA data of species of Leptolycus in Puerto Rico. Utilizing the same biogeography-based rationale with C. tortolensis in Tortola and combining it with our findings and observations on T. viensis, we were able to predict the overall phenotype of other species of Leptolycus and Cessator also present in Puerto Rico, which was later corroborated by our DNA-based analyses. Conclusion Our study is the first to provide life-stage associations of Leptolycini based on the use of biogeography and DNA-based data. Whilst the main goal of our study was to provide such associations, these could not have been done without the extensive clarifications in the taxonomy of the group in the Puerto Rican bank. The taxonomic work herein presented resulted in important changes in the classification of the group, with consequences that go beyond the simple description of new taxa in the tribe. We provided a review of the Leptolycini taxa in the Puerto Rican bank, a key to the genera and all species of adults of the region, as well as an extensive debate on the intraspecific variation in members of the group, resulting in the descriptions of six new species, a new genus and the synonymy of one genus, one subgenus and one species. As consequences of our life-stage associations, we also provided the first illustrations, diagnoses and identification keys of the immature forms and extreme paedomorphic females of the Leptolycini. Even though our results significantly expanded the knowledge of the group and provided evidence of extreme color polymorphism in both the larvae and adults in Leptolycus, several questions remain unresolved. Among these, we would like to further investigate the intraspecific variability of the several color morphs found in L. heterocornis and to obtain DNA grade specimens from all the morphs of this species to compare and confront them with our proposed morphological species delimitations. We also would like to build a COI library of all Leptolycini adults and larvae of the Puerto Rican bank, to provide a definitive match of Dracolycus and their respective hypothesized immature forms and identify the immature forms and extreme paedomorphic females of all other species and genera in the region. For these, the identification of further material in scientific collections and further collecting efforts are necessary, and we hope that with the publication of this study we can capture the attention of entomologists and collectors. At last, we hope to have provided another example of the importance of natural history collections in taxonomic, systematic, and evolutionary works. While the utilization of modern and more sophisticated DNA-based techniques significantly increases our taxonomic and systematic knowledge of the groups, without the historical material of both adult males and larvae deposited in collections, these conclusions, and further advances in the poorly understood phenomenon of paedomorphosis is would not have been possible. Acknowledgments The collaborators across the world are too numerous to list here, and we are grateful for all their efforts. VSF is grateful to the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) of Brazil for his PhD scholarship (process 202559/2015-7) and to the Ernst Mayr Travel Grants in Animal Systematics; the Systematics, Evolution, and Biodiversity (SysEB) Student Research Travel Award from the Entomological Society of America and the several Montana State University Graduate School and College of Agriculture travel awards that allowed the visit to several collections and the field expedition to Puerto Rico; to the Idea Wild Project for providing resources that allowed the acquisition of part of the photographic equipment used in this project; to Erich L. Spiessberger for his assistance and pleasant company and friendship during VSF fieldwork in Puerto Rico. Authors are grateful to the respective curators and collections managers of each collection who provided material for this study and to the many persons who collaborated either with specimens or other valuable information, especially to Sergey Kazantsev (Insect Centre) and Matt Gimmel (Santa Barbara Museum of Natural History) for tips for collecting adult male Leptolycini in the El Verde Field Station and to Donald Yee (University of Southern Mississippi) for the donation of several critical larvae specimens that were used in this study. We thank the late Richard S. Miller for the critical donation of his Lycidae collection to the WIBF and for his valuable suggestions and discussions during the development of this and several other studies with other soft-bodied Elateroidea. Several larvae specimens were collected by students funded by a grant to Donald Yee from the National Science Foundation (DEB-1806122). Research in the El Verde Field Station was possible due to the support grants BSR-8811902, DEB 9411973, DEB 0080538, DEB 0218039, DEB 0620910, DEB 0963447, and DEB-1546686 from National Science Foundation to the Institute for Tropical Ecosystem Studies, University of Puerto Rico, and to the International Institute of Tropical Forestry USDA Forest Service, as part of the Luquillo Long-Term Ecological Research Program, with additional support from the U.S. Forest Service (Dept. of Agriculture) and the University of Puerto Rico. This is a contribution of the Montana Agricultural Experiment Station. Author Contributions VSF and MAI conceived the study, discussed results, interpreted results, and revised the manuscripts versions. VSF examined all specimens, did all dissections, extracted, sequenced, processed, and analyzed all DNA samples and performed phylogenetic analyses, led all the discussions, made all illustrations, wrote the manuscript. Both authors agreed with the final submitted version of this manuscript. Version of Record, first published online May 17, 2022 with fixed content and layout in compliance with Art. 8.1.3.2 ICZN References Cited Beatty , H. A . 1944 . Fauna of St. Croix, Virgin Islands . J. Agric. Univ. P.R . 28 : 103 – 185 . Google Scholar OpenURL Placeholder Text WorldCat Bocak , L . 2001 . New taxa of Leptolycus (Coleoptera: Lycidae) from the Dominican Republic . Folia Heyrovkyana . 9 : 203 – 210 . Google Scholar OpenURL Placeholder Text WorldCat Bocak , L . 2018 . Dexoris ruzzieri sp. nov. (Coleoptera: Lycidae), a new net-winged beetle from Ivory Coast . Zootaxa . 4377 ( 1 ): 138 – 142 . doi:10.11646/zootaxa.4377.1.9 Google Scholar Crossref Search ADS PubMed WorldCat Bocak , L. , and M. Bocákova. 1990 . Revision of the suprageneric classification of the family Lycidae (Coleoptera) . Pol. Pismo Entomo . 59 : 623 – 676 . Google Scholar OpenURL Placeholder Text WorldCat Bocak , L. , and M. Bocakova. 2008 . Phylogeny and classification of the family Lycidae (Insecta: Coleoptera) . Ann. Zool . 58 : 695 – 720 . Google Scholar Crossref Search ADS WorldCat Bocak , L. , and K. Matsuda. 2003 . Review of the immature stages of the family Lycidae (Insecta: Coleoptera) . J. Nat. Hist . 37 : 1463 – 1507 . Google Scholar Crossref Search ADS WorldCat Bocak , L. , M. Bocakova, T. Hunt, and A. P. Vogler. 2008 . Multiple ancient origins of neoteny in Lycidae (Coleoptera): consequences for ecology and macroevolution . Proc. Royal Soc. B . 275 : 2015 – 2023 . Google Scholar Crossref Search ADS WorldCat Bocakova , M . 2003 . Revision of the tribe Calopterini (Coleoptera, Lycidae) . Neotrop. Fauna Environ . 38 : 207 – 234 . Google Scholar Crossref Search ADS WorldCat Bocakova , M . 2005 . Phylogeny and classification of the tribe Calopterini (Coleoptera, Lycidae) . Insect Syst. Evol . 35 : 437 – 447 . Google Scholar Crossref Search ADS WorldCat Bocakova , M . 2014 . Lolodorfus, a new genus of net-winged beetles (Coleoptera: Lycidae: Dexorinae) from Cameroon . Zootaxa 3811 : 374 – 380 . Google Scholar Crossref Search ADS WorldCat Bocakova , M. , L. Bocak, T. Hunt, M. Teravainen, and A. P. Vogler. 2007 . Molecular phylogenetics of Elateriformia (Coleoptera): evolution of bioluminescence and neoteny . Cladistics 23 : 477 – 496 . Google Scholar Crossref Search ADS WorldCat Bocek , M. , L. Fancello, M. Motyka, M. Bocakova, and L. Bocak. 2018 . The molecular phylogeny of Omalisidae (Coleoptera) defines the family limits and demonstrates low dispersal propensity and ancient vicariance patterns . Syst. Entomol . 43 : 250 – 261 . doi:10.1111/syen.12271 Google Scholar Crossref Search ADS WorldCat Bocek , M. , M. Motyka, D. Kusy, and L. Bocak. 2019 . Genomic and mitochondrial data identify different species boundaries in aposematically polymorphic eniclases net-winged beetles (Coleoptera: Lycidae) . Insects . 10 : 295 . Google Scholar Crossref Search ADS WorldCat Blackwelder , R. E . 1945 . Checklist of the coleopterous insects of Mexico, Central America, the West Indies, and South America, pt. 3 . Bull. Am. Mus. Nat. Hist . i–iv : 343 – 550 . Google Scholar OpenURL Placeholder Text WorldCat Brown , S. , Lugo , A.E., Silander , S., and Liegel , L. 1983 . Research history and communities in the Luquillo experimental forest . U.S. Department of Agriculture, Forest Service, General Technical Report SO-44, New Orleans, Louisiana : Southern Forest Experiment Station . Google Scholar Google Preview OpenURL Placeholder Text WorldCat COPAC Cicero , J. M . 1988 . Ontophylogenetics of Cantharoid Larviforms (Coleoptera: Cantharoidea) . Coleopt. Bull . 42 : 105 – 151 . Google Scholar OpenURL Placeholder Text WorldCat Costa , C. , S.A. Casari-Chen, and S.A. Vanin. 1988 . Larvas de Coleoptera do Brasil. Museu de Zoologia, Universidade de São Paulo . 282 + 65 plates pp. https://doi.org/10.5962/bhl.title.100233 Crowson , R. A . 1972 . A review of the classification of Cantharoidea (Coleoptera), with definition of two new families: Cneoglossidae and Omethidae . Revista de la Universidad de Madrid 21 : 35 – 77 . Google Scholar OpenURL Placeholder Text WorldCat Etzler , F. E. , J. Huether, P. J. Johnson, F. W. Skillman , Jr., and M. A. Ivie. 2013 . Use of morphology and CO1 barcoding to test the validity of Tetraopes huetheri Skillman (Coleoptera: Cerambycidae) . Coleop. Bull . 67 : 46 – 49 . Google Scholar Crossref Search ADS WorldCat Etzler , F. E. , K. W. Wanner, A. Morales-Rodriguez, and M. A. Ivie. 2014 . DNA barcoding to improve the species-level management of wireworms (Coleoptera: Elateridae) . J. Econ. Entomol . 107 : 1476 – 1485 . Google Scholar Crossref Search ADS PubMed WorldCat EVFS [El Verde Field Station website]. 2022 . <https://luq.lter.network/el-verde-field-station-el-verde> Last access April 12, 2022 . Ferreira , V. S . 2020 . Revision of Acroleptus Bourgeois, 1886 and descriptions of new Aporrhipis species (Lycidae, Calopterini, Acroleptina) . J. Nat. Hist . 53 : 2739 – 2756 . Google Scholar Crossref Search ADS WorldCat Ferreira , V. S. , and C. Costa. 2015 . A description of the larva of Metapteron xanthomelas (Lucas, 1857) from the restinga forest of Southeastern Brazil (Coleoptera: Lycidae, Calopterini) . Zootaxa 3915 : 295 – 300 . Google Scholar Crossref Search ADS PubMed WorldCat Ferreira , V. S. , and C. Costa. 2017 . An online database of the immatures of Coleoptera (Arthopoda, Insecta) described from Brazil . Bio. Data. J . 5 : e12252 . Google Scholar OpenURL Placeholder Text WorldCat Ferreira , V. S. , and M. A. Ivie. 2016 . Redescription of Cephalolycus Pic, 1926 (Coleoptera: Elateroidea: Lycidae) and a discussion on its taxonomic position . Col. Bull . 70 : 663 – 666 . Google Scholar Crossref Search ADS WorldCat Ferreira , V. S. , and M. A. Ivie. 2017 . The first fossil species of the extant genus Cessator Kazantsev (Coleoptera: Lycidae): a new leptolycini from Dominican Amber . Coleopt. Bull . 71 : 57 – 60 . Google Scholar Crossref Search ADS WorldCat Ferreira , V. S. , and M. A. Ivie. 2018 . A revision of Lycinella Gorham, 1884 with the description of six new species (Coleoptera, Lycidae, Calopterini) . ZooKeys . 792 : 69 – 89 . Google Scholar Crossref Search ADS WorldCat Ferreira , V. S. , and L. F. L. Silveira. 2020 . A new suspected paedomorphic genus of net-winged beetles from the Atlantic Rainforest (Coleoptera, Elateroidea, Lycidae) . Pap. Avulsos Zool . 60 ( special ): e202060(s.i.) .35:1–6. https://doi.org/10.11606/1807-0205/2020.60.special-issue.35 Google Scholar OpenURL Placeholder Text WorldCat Ferreira , V. S. , O. Keller, M. A. Branham, and M. A. Ivie. 2019 . Molecular data support the placement of the enigmatic Cheguevaria as a subfamily of Lampyridae (Insecta: Coleoptera) . Zool. J. Linn. Soc . 187 : 1253 – 1258 . Google Scholar Crossref Search ADS WorldCat Ferreira , V. S. , O. Keller, and M. A. Branham. 2020 . Multilocus phylogeny support the nonbioluminescent firefly Chespirito as a new subfamily in the Lampyridae (Coleoptera: Elateroidea) . Insect Syst. Div . 4 : 1 – 13 . Google Scholar OpenURL Placeholder Text WorldCat Ferreira , V. S. , A. Solodvnikov, M. A. Ivie, and R. Kundrata. 2022 . Dominican Amber Net-Winged Beetles suggest stable Paleoenvironment as a driver for Conserved Morphology in a Paedomorphic lineage . Sci. Rep . 12 : 5820 : 1–10. doi:10.1038/s41598-022-09867-6 Google Scholar Crossref Search ADS PubMed WorldCat Fitch , W. M. , and E. Margoliash. 1967 . Construction of phylogenetic trees . Science 155 : 279 – 284 . Google Scholar Crossref Search ADS PubMed WorldCat Folmer , O. , M. Black, W. Hoeh, R. Lutz, and R. Vrijenhoek. 1994 . DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates . Mol. Mar. Biol. Biotech , 3 : 294 – 299 . Google Scholar OpenURL Placeholder Text WorldCat ICZN . 1999 . International Code of Zoological Nomenclature, Fourth Edition, adopted by the International Union of Biological Sciences . International Trust for Zoological Nomenclature , London , xxix + 306 pp. Available from http://www.iczn.org/iczn/index.jsp. Google Scholar Google Preview OpenURL Placeholder Text WorldCat COPAC Jiruskova , A. , and L. Bocak. 2015 . Species delimitation in Cautires (Coleoptera: Lycidae) from Peninsular Malaysia using DNA data and morphology . Ann. Zool . 65 : 239 – 248 . Google Scholar Crossref Search ADS WorldCat Jukes , T.H. , and C.R. Cantor. 1969 . Evolution of protein molecules. pp. 21 – 132 . In H.N. Munro (ed.), Mammalian Protein Metabolism . Academic Press , New York . Google Scholar Crossref Search ADS Google Preview WorldCat COPAC Katoh , K. , K. Misawa, K. I. Kuma, and T. Miyata. 2002 . MAFFT: a novel method for rapid multiple sequence alignment based on fast Fourier transform . Nucl. Acids Res . 30 : 3059 – 3066 . Google Scholar Crossref Search ADS WorldCat Kazantsev , S. V . 2005 . Morphology of Lycidae with some considerations on evolution of the Coleoptera . Elytron . 17 : 49 – 226 . Google Scholar OpenURL Placeholder Text WorldCat Kazantsev , S. V . 2009 . Leptolycini of Puerto Rico (Coleoptera: Lycidae) . Russ. Entom. J . 18 : 87 – 95 . Google Scholar OpenURL Placeholder Text WorldCat Kazantsev , S. V . 2012 . A new lycid genus from the Dominican Amber (Insecta, Coleoptera, Lycidae, Leptolycinae, Leptolycini) . Psyche . 982141 : 1 – 3 . doi:10.1155/2012/982141 Google Scholar OpenURL Placeholder Text WorldCat Kazantsev , S. V . 2013 . New and little-known taxa of ‘neotenic’ Lycidae (Coleoptera), with discussion of their phylogeny . Russ. Entom. J . 22 : 9 – 31 . Google Scholar OpenURL Placeholder Text WorldCat Kazantsev , S. V . 2017 . New leptolycines from Ecuador and Peru (Coleoptera: Lycidae) . Russ. Entom. J . 26 : 127 – 146 . Google Scholar Crossref Search ADS WorldCat Kazantsev , S. V. and A. A. Zaitsev. 2008 . Description of larval and imaginal stages of new species from the genera Pseudacroleptus Pic, 1911 and Ceratoprion Gorham, 1880 (Coleoptera: Lycidae: Leptolycinae) . Russ. Entomol. J . 17 ( 3 ): 283 – 292 . Google Scholar OpenURL Placeholder Text WorldCat Keller , O. , and M. Branham. 2021 . A catalog of the Lampyridae (Coleoptera) of the West Indies . Zootaxa 4940 : 53 – 105 . Google Scholar OpenURL Placeholder Text WorldCat Kimura , M . 1980 . A simple method for estimating evolutionary rates of base substitutions through comparative studies of nucleotide sequences . J. Mol. Evol . 16 : 111 – 120 . Google Scholar Crossref Search ADS PubMed WorldCat Kleine , R . 1933 . Lycidae Pars 128 . In W. Junk, S. Schenkling (eds.), Coleopterorum Catalogueus auspiciis et auxilio . Berlin , 145 pp. Google Scholar Google Preview OpenURL Placeholder Text WorldCat COPAC Kumar , S. , G. Stecher, M. Li, C. Knyaz, and K. Tamura. 2018 . MEGA X: Molecular Evolutionary Genetics Analysis across computing platforms . Mol. Biol. Evol . 35 : 1547 – 1549 . Google Scholar Crossref Search ADS PubMed WorldCat Kundrata , R. , and L. Bocak. 2007 . A revision of Euanoma and Pseudeuanoma (Coleoptera: Drilidae) . Ann. Zool . 57 ( 3 ): 427 – 441 . Google Scholar OpenURL Placeholder Text WorldCat Kundrata , R. , and L. Bocak. 2019 . Molecular phylogeny reveals the gradual evolutionary transition to soft-bodiedness in click-beetles and identifies Sub-Saharan Africa as a cradle of diversity for Drilini . Zool. J. Linn. Soc . 187 : 413 – 452 . Google Scholar Crossref Search ADS WorldCat Kundrata , R. , M. Bocakova, and L. Bocak. 2014 . The comprehensive phylogeny of the superfamily Elateroidea (Coleoptera: Elateriformia) . Mol. Phylogenet. Evol . 76 : 162 – 171 . Google Scholar Crossref Search ADS PubMed WorldCat Kundrata , R. , S. M. Blank, A. S. Prosvirov, E. Sormova, M. L. Gimmel, D. Vondráček, and K. Kramp. 2019 . One less mystery in Coleoptera systematics: the position of Cydistinae (Elateriformia incertae sedis) resolved by multigene phylogenetic analysis . Zool. J. Linn. Soc . 187 : 1259 – 1277 . Google Scholar Crossref Search ADS WorldCat Kusy , D. , M. Motyka, M. Bocek, A. P. Vogler, and L. Bocak. 2018a . Genome sequences identify three families of Coleoptera as morphologically derived click beetles (Elateridae) . Sci. Rep . 8 : 17084 . Google Scholar Crossref Search ADS WorldCat Kusy , D. , K. Sklenarova, and L. Bocak. 2018b . The effectiveness of DNA-based delimitation in Synchonnus net-winged beetles (Coleoptera: Lycidae) assessed, and description of 11 new species . Austral Ent . 57 : 25 – 39 . Google Scholar Crossref Search ADS WorldCat Kusy , D. , M. Motyka, M. Bocek, M. Masek, and L. Bocak. 2019 . Phylogenomic analysis resolves the relationships among net-winged beetles (Coleoptera: Lycidae) and reveals the parallel evolution of morphological traits . Syst. Ent . 44 : 911 – 925 . Google Scholar Crossref Search ADS WorldCat Lawrence , J. F. , A. Slipinski, A. E. Seago, M. K. Thayer, A. F. Newton, and A. E. Marvaldi. 2011 . Phylogeny of the Coleoptera based on morphological characters of adults and larvae . Ann. Zool . 61 : 1 – 217 . Google Scholar Crossref Search ADS WorldCat Leng , C. W. , and A. J. Mutchler. 1922 . Article VIII.- The Lycidae, Lampyridae and Cantharidae (Telephoridae) of the West Indies . Bull. Am. Mus. Nat. Hist . 46 : 413 – 499 . Google Scholar OpenURL Placeholder Text WorldCat Levkanikova , Z. , and L. Bocak. 2009 . Identification of net-winged beetle larvae (Coleoptera: Lycidae) using three mtDNA fragments: a comparison of their utility . Syst. Ent . 34 : 210 – 222 . Google Scholar Crossref Search ADS WorldCat Li , Y. , H. Pang, and L. Bocak. 2018a . A review of the neotenic genus Atelius Waterhouse, 1878 from China (Coleoptera: Lycidae) . Ann. Zool . 68 : 351 – 356 . Google Scholar Crossref Search ADS WorldCat Li , Y. , N. Gunter, H. Pang, and L. Bocak. 2018b . DNA-based species delimitation separates highly divergent populations within morphologically coherent clades of poorly dispersing beetles . Zool. J. Linn. Soc . 175 : 59 – 72 . Google Scholar Crossref Search ADS WorldCat Linnaeus , C . 1763 . Amoenitates Academicae. VI , pp. 508 . Google Scholar Google Preview OpenURL Placeholder Text WorldCat COPAC Martin , G. J. , M. A. Branham, M. F. Whiting, and S. M. Bybee. 2017 . Total evidence phylogeny and the evolution of adult bioluminescence in fireflies (Coleoptera: Lampyridae) . Mol. Phyl. Evol . 564 – 575 . Google Scholar OpenURL Placeholder Text WorldCat Martin , G. J. , K. Stanger-Hall, M. A. Branham, L. F. L. Silveira, S. E. Lower, D. W. Hall, X. Li, A. R. Lemmon, E. M. Lemmon, and S. M. Bybee. 2019 . Higher-level phylogeny and reclassification of Lampyridae (Coleoptera: Elateroidea) . Inse. Syst. Div . 3 ( 6 ): 11; 1 – 15 . Google Scholar OpenURL Placeholder Text WorldCat Masek , M. , and L. Bocak. 2014 . The taxonomy and diversity of Platerodrilus (Coleoptera, Lycidae) inferred from molecular data and morphology of adults and larvae . ZooKeys 426 : 29 – 63 . Google Scholar OpenURL Placeholder Text WorldCat Masek , M. , M. A. Ivie, V. Palata, and L. Bocak. 2014 . Molecular phylogeny and classifi cation of Lyropaeini (Coleoptera: Lycidae) with description of larvae and new species of Lyropaeus . Raff. Bull. Zool . 62 : 136 – 145 . Google Scholar OpenURL Placeholder Text WorldCat Masek , M. , M. Motyka, D. Kusy, M. Bocek, Y. Li, and L. Bocak. 2018 . Molecular phylogeny, diversity and zoogeography of net-winged beetles (Coleoptera: Lycidae) . Insects . 9 : 154 . Google Scholar Crossref Search ADS WorldCat McDowell , W. H. , F. N. Scatena, W. B. Waide, N. Brokaw, G. R. Camilo, A. P. Covich, T. A. Crowl, G. González, E. A. Greathouse, P. Klawinski, et al. . 2012 . Geographic and ecological setting of the Luquillo Mountains. pp. 72 – 163 . In N. Brokaw, T. A. Crowl, A. E. Lugo, W. H. McDowell, F. N. Scatena, R. B. Waide, M. R. Willig (eds.), A Caribbean forest tapestry: the multidimensional nature of disturbance and response . Oxford University Press , New York . Google Scholar Crossref Search ADS Google Preview WorldCat COPAC McMahon , D. P. , and A. Hayward. 2016 . Why grow up? A perspective on insect strategies to avoid metamorphosis . Ecol. Entomo . 41 : 505 – 515 . Google Scholar Crossref Search ADS WorldCat McNamara , K. J . 2012 . Heterochrony: the evolution of development . Evol.: Educ. Outreach . 5 : 203 – 218 . Google Scholar OpenURL Placeholder Text WorldCat Mertins , J. W . 1981 . Life history and morphology of the odd beetle, Thylodrias contractus . Ann. Entomol. Soc. Am . 74 : 576 – 581 . Google Scholar Crossref Search ADS WorldCat Miller , R. S . 1991 . A revision of Leptolycini (Coleoptera: Lycidae) with a discussion of paedomorphosis . PhD Dissertation. The Ohio State University . 1 – 403 pp (unpublished). Google Scholar Google Preview OpenURL Placeholder Text WorldCat COPAC Miskimen , G. W. , and R. M. Bond. 1970 . The insect fauna of St. Croix, U. S. Virgin Islands. Scientific Survey of Porto Rico and the Virgin Islands . N. Y. Acad. Sci . 13 : 1 – 117 . Google Scholar OpenURL Placeholder Text WorldCat Mjöberg , E . 1925 . The mystery of the so called ‘Trilobite’ or ‘Perty’s larvae’ defi nitely solved . Psyche . 22 : 119 – 153 . Google Scholar OpenURL Placeholder Text WorldCat Perotti , M. A. , D. K. Young, and H. R. Braig. 2016 . The ghost sex-life of the paedogenetic beetle Micromalthus debilis . Sci. Rep . 6 : 27364 . Google Scholar Crossref Search ADS PubMed WorldCat Ratnasingham , S. , and P. D. N. Hebert. 2013 . A DNA-based registry for all animal species: the Barcode Index Number (BIN) system . PLoS One 8 : 1 – 16 . doi:10.1371/journal.pone.0066213 Google Scholar Crossref Search ADS WorldCat Rattu , R . 2016 . Contributo alla conoscenza delle femmine delle specie sarde di Cebrio (Coleoptera, Elateridae, Elaterinae, Cebrionini). Annali Mus. civ. St. nat. “G. Doria” Genova 108 : 235 – 248 . Rattu , R . 2020 . Morphological, geonemic and ecological notes on Cebrio (Tibesia) mediatlantis mediatlantis Kocher, 1952, and description of the female (Coleoptera, Elateridae, Elaterinae, Cebrionini) . Boll. Mus. Civico Storia Nat. Venezia . 71 : 51 – 59 . Google Scholar OpenURL Placeholder Text WorldCat Raupach , M. J. , K. Hannig, J. Morinière, and L. Hendrich. 2016 . A DNA barcode library for ground beetles (Insecta, Coleoptera, Carabidae) of Germany: The genus Bembidion Latreille, 1802 and allied taxa . ZooKeys 592 : 121 – 141 . Google Scholar Crossref Search ADS WorldCat Reilly , S. M. , E. O. Wiley, and D. J. Meinhardt. 1997 . An integrative approach to heterochrony: the distinction between interspecific and intraspecific phenomena . Biol. J. Linn. Soc . 60 : 119 – 143 . Google Scholar Crossref Search ADS WorldCat Reiss , R. A. , D. P. Schwert, and A. C. Ashworth. 1995 . Field preservation of Coleoptera for molecular genetic analyses . Env. Ent . 24 : 716 – 719 . Google Scholar Crossref Search ADS WorldCat Roza , A. S. , H. Y. S. Quintino, J. R. M. Mermudes, and L. F. L. Silveira. 2017 . Akamboja gen. nov., a new genus of railroad-worm beetle endemic to the Atlantic Rainforest, with five new species (Coleoptera: Phengodidae, Mastinocerinae) . Zootaxa 4306 : 501 – 523 . Google Scholar Crossref Search ADS WorldCat Schowalter , T. D. , M. R. Willig, and S. J. Presley. 2014 . Canopy arthropod responses to experimental canopy opening and debris deposition in a tropical rainforest subject to hurricanes . For. Ecol. Man . 332 : 93 – 102 . Google Scholar Crossref Search ADS WorldCat Sikes , D. S. , and L. J. Mullen. 2021 . Phylogeny and evolution of large body size in the rove beetle genus Phlaeopterus Motschulsky, 1853 (Coleoptera: Staphylinidae: Omaliinae: Anthophagini) . Arthr. Syst. Phyl . 70 : 75 – 98 . Google Scholar OpenURL Placeholder Text WorldCat Srivathsan , A. , B. Baloğlu, and W. Wang, et al. . 2018 . A MinION™-based pipeline for fast and cost-effective DNA barcoding . Mol. Ecol. Res . 18 : 1035 – 1049 . Google Scholar Crossref Search ADS WorldCat Srivathsan , A. , E. Hartop, and J. Puniamoorthy, et al. . 2019 . Rapid, large-scale species discovery in hyperdiverse taxa using 1D MinION sequencing . BMC Biol . 17 : 96 . Google Scholar Crossref Search ADS PubMed WorldCat Telfer , A. C. , M. R. Young, J. Quinn, K. Perez, C. N. Sobel, Sones, et al. . 2015 . Biodiversity inventories in high gear: DNA barcoding facilitates a rapid biotic survey of a temperate nature reserve . Biod. Data. J . 3 : e6313: 1 – 176 . doi:10.3897/BDJ.3.e6313 Google Scholar OpenURL Placeholder Text WorldCat Trifinopoulos , J. , L. T. Nguyen, A. von Haeseler, and B. Q. Minh. 2016 . IQ-TREE web server: fast and accurate phylogenetic trees under maximum likelihood . Nuc. Acids Res . 44 : W232 – W235 . Google Scholar Crossref Search ADS WorldCat Vea , I. M. , and C. Minakuchi. 2021 . Atypical insects: molecular mechanisms of unusual life history strategies . Curr. Opi. Insect Sci 43 : 46 – 53 . Google Scholar Crossref Search ADS WorldCat Wong , A. T. C . 1996 . A new species of neotenous beetle, Duliticola hoiseni (Insecta: Coleoptera: Cantharoidea: Lycidae) from peninsular Malaysia and Singapore . Raff. Bull. Zool . 44 : 173 – 187 . Google Scholar OpenURL Placeholder Text WorldCat Wu , R. J. C . 1997 . Secrets of a lost world: Dominican Amber and its inclusions . Privately published , Santo Domingo, Dominican Republic . Google Scholar Google Preview OpenURL Placeholder Text WorldCat COPAC Zapata de La Vega , J. S. , A. Sánchez-Ruiz, and M. A. López. 2020 . Nueva aportación al conocimiento del género Cebrio Olivier,1790. Descripción de la hembra de Cebrio (Tibesia) rozasi Cobos, 1985 (Coleoptera: Elateridae: Elaterinae: Cebrionini) . Bol. SAE . 30 : 69 – 74 . Google Scholar OpenURL Placeholder Text WorldCat de Zayas , F. . 1988 . Entomofauna Cubana. Orden Coleoptera. Separata. Descripcion de Nuevas Especies . Editorial Cientifico-Tecnica . 212 pp. Google Scholar Google Preview OpenURL Placeholder Text WorldCat COPAC Zhai , Q. , G. Xue, and M. Li. 2017 . DNA barcoding-based sexual association of Sovia lucasii and S. lii (Lepidoptera: Hesperiidae), with description of a new subspecies . PLoS One 12 : e0183847 . Google Scholar Crossref Search ADS PubMed WorldCat Zilberman , B . 2020 . Phylogenetic analysis of the genus Corotoca, with description of a new genus and species from Brazil (Coleoptera, Staphylinidae, Aleocharinae) . Insec. Syst. Evol . 52 : 201 – 245 . Google Scholar Crossref Search ADS WorldCat © The Author(s) 2022. Published by Oxford University Press on behalf of Entomological Society of America. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com. 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TI - Lessons From a Museum's Cabinet: DNA Barcoding and Collections-Based Life Stage Associations Reveals a Hidden Diversity in the Puerto Rican Bank Paedomorphic Lycidae (Coleoptera: Elateroidea: Leptolycini)
JF - Insect Systematics and Diversity
DO - 10.1093/isd/ixac006
DA - 2022-05-01
UR - https://www.deepdyve.com/lp/oxford-university-press/lessons-from-a-museum-s-cabinet-dna-barcoding-and-collections-based-a6ofwnpzGD
VL - 6
IS - 3
DP - DeepDyve
ER -