Research Article |
Corresponding author: Emmanuel Arriaga-Varela ( arriagavarelae@natur.cuni.cz ) Academic editor: Mariano Michat
© 2017 Emmanuel Arriaga-Varela, Matthias Seidel, Albert Deler-Hernández, Viktor Senderov, Martin Fikáček.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Arriaga-Varela E, Seidel M, Deler-Hernández A, Senderov V, Fikáček M (2017) A review of the Cercyon Leach (Coleoptera, Hydrophilidae, Sphaeridiinae) of the Greater Antilles. ZooKeys 681: 39-93. https://doi.org/10.3897/zookeys.681.12522
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The representatives of the genus Cercyon Leach occurring in the Greater Antilles are reviewed. Ten species are recorded, of which five are described here as new: C. gimmeli sp. n. (Dominican Republic), C. armatipenis sp. n. (Dominican Republic), C. taino sp. n. (Dominican Republic), C. sklodowskae sp. n. (Jamaica) and C. spiniventris sp. n. (Dominican Republic). Diagnoses and detailed distributional data are also provided for C. floridanus Horn, 1890 (distributed in southeastern United States of America and Cayman Islands), C. insularis Chevrolat, 1863 (endemic to the Antilles) C. praetextatus (Say, 1825) (widely distributed in the New World incl. Greater Antilles), C. quisquilius (Linnaeus, 1761) (an adventive species of Paleartic origin) and C. nigriceps (Marshall, 1802) (an adventive species probably of Oriental origin). Cercyon armatipenis, C. gimmeli, C. taino form a group of closely related species only distinguishable by male genitalia and DNA sequences. A key to the Great Antillean Cercyon is provided and important diagnostic characters are illustrated. The larvae of C. insularis and C. taino were associated with adults using COI barcode sequences, illustrated and diagnosed. Full occurrence data, additional images and COI barcode sequences were submitted to open access on-line depositories in an effort to provide access to complete data.
Megasternini , morphology, taxonomy, new species, Caribbean, COI, DNA barcode, larva, biodiversity informatics
Until very recently, water scavenger beetles (Hydrophilidae) from the Greater Antilles in the Caribbean Region were largely neglected and systematic or faunistic studies were scarce. For instance, 46 of 56 hydrophilid species recorded from Cuba were described in the 18th and 19th centuries (
Six species of Cercyon have been recorded from the Greater Antilles: the Cuban-endemic C. insularis Chevrolat whose identity has remained unclear (
Examined specimens and depositories. A total of 848 specimens of Cercyon from the Greater Antilles were examined, including the type specimens of Cercyon insularis Chevrolat and C. variegatus Sharp. Label data are only reproduced verbatim for type specimens; each individual label is separated by double slash “//”; notes on the label data or additional information are written between square brackets []. All holotypes are marked with red label bearing the following text: “HOLOTYPE, Cercyon [name of the species] sp. n., Arriaga-Varela, Seidel, Deler-Hernández and Fikáček des. 2016“. All paratype specimens are marked by yellow label bearing the following text: “PARATYPE, Cercyon [name of the species] sp. n., Arriaga-Varela, Seidel, Deler-Hernández and Fikáček des. 2016”. A georeferenced dataset of the studied specimens is available as Excel spreadsheet in Suppl. material
The examined specimens are deposited in the following collections:
BCPC Bruno Clarkson private collection, Rio de Janeiro, Brazil;
SBPC Stewart Peck Personal Collection, Ottawa, Canada;
Morphological studies. Specimens were dissected, with genitalia embedded in a drop of alcohol-soluble Euparal resin on a piece of glass glued to a small piece of cardboard attached below the respective specimen. All species are diagnosed and illustrated, and new species are described in detail.
Habitus photographs were taken using a Canon D-550 digital camera with attached Canon MP-E65mm f/2.8 1–5 macro lens. Pictures of genitalia were taken using a Canon D1100 digital camera attached to an Olympus BX41 compound microscope; pictures of different focus were combined in Helicon Focus software. Scanning electron micrographs were taken using Hitachi S-3700N environmental electron microscope at the Department of Paleontology, National Museum in Prague. Pictures used for plates were adapted in Adobe Photoshop CS6. All original pictures including additional views not presented in this paper are published and freely available on Flickr in order to serve for further morphological studies.
DNA barcoding. Most of the examined specimens were collected during recent expeditions to Cuba, Dominican Republic and Puerto Rico. Samples were preserved in 96% ethanol and stored at -20 °C. DNA was extracted from complete specimens using a QiaGen Blood and Tissue DNA extraction kit following the manufacturer's instructions. The highly variable 5’ region of the mitochondrial cytochrome c oxidase subunit I gene (COI) was amplified using LCO1490 (5'-GGTCAACAAATCATAAAGATATTGG-3') and HCO2198 (5'-TAAACTTCAGGGTGACCAAAAAATCA-3') primers (Folmer et al. 1994). Each 10 µl PCR reaction contained 6.7 µl H2O, 0.4 µl of MgCl2 (25 mM), 0.2 µl of dNTPs (10 mM), 0.3 µl of each forward and reverse primer (10 µM), 0.1 µl of Taq polymerase (5 u/µl), 1.0 µl of 10x Taq buffer, and 1.0 µl of DNA template. The PCR conditions consisted of 3 min at 94 °C + 35 cycles of 30 s at 94 °C, 45 s at 48 °C and 1 min at 72 °C + 8 min at 72 °C. 5 µl of each PCR product were purified by adding 0.5µl (20 u) Exonuclease I (Exo1) and 1µl (1 u) Thermosensitive Alkaline Phosphatase (FastAP) (Thermo Fisher Scientific) and incubating the mixture for 15 min at 37°C, followed by 15 min at 80°C. The Sanger sequencing was performed by BIOCEV (Vestec, Czech Republic) on a capillary DNA sequencer. Sequences were edited with Geneious 9.1.4. We did not attempt DNA extraction and sequencing of old museum specimens, which is why we only provide sequences for six of the ten species occurring in Greater Antilles.
Analyses of molecular data. In order to identify the larvae collected along with adult specimens, a maximum likelihood analysis of obtained COI sequences was performed. We combined the newly generated sequences of freshly collected adults and larvae, and combined them with additional sequences of two introduced species (C. quisquilius and C. nigriceps from Europe and Canada) from the Barcode of Life Data Systems (BOLD; http://www.boldsystems.org). Sequences were aligned using the ClustalW algorythm in Geneious 9.1.4. The final alignment had a length of 610 bp and was tested for the best nucleotide substitution model using MEGA7 (
This taxonomic paper includes only a part of the data accumulated in the course of our study. Part of the primary data (e.g., unedited photographs, the complete set of unedited SEM micrographs, DNA sequences, spreadsheet-fomatted species distribution data) are not included here. To make all primary data accessible, we deposited them to open access on-line depositories as specified below. For more details about biodiversity data publishing, see e.g. the policies and guidelines implemented for Pensoft Publishers (
Complete primary data. Complete primary data were submitted as a .zip file to the Zenodo depository (https://zenodo.org/) under the doi 10.5281/zenodo.580260.
Species distribution data. The distribution data on all specimens examined are presented in unstructured text format directly in the paper. The conversion of these data into a structured, computable format (as XML, so called parsing) is difficult, and no algorithm exists for parsing occurrence records (see
The text-formatted distribution data published here are, however, based on a structured Excel spreadsheet following the Darwin Core (henceforth DwC) format for biodiversity data described by
The Publication of distribution data to GBIF is possible through the institution or organization, which is a member or partner of GBIF (direct submissions from individual users are not possible) using the Integrated Publishing Toolkit (IPT). IPT allows to upload the distribution data from the DwC-formatted Excel spreadsheet, specify the metadata about the dataset, and publish the data to the GBIF portal. We submitted our data through Pensoft as an organization associated with GBIF using the Pensoft IPT Data Hosting Centre (http://ipt.pensoft.net/).
DNA data and voucher information. The cytochrome oxidase I barcode sequences and the data about the voucher specimens were submitted to the Barcode of Life Data Systems (BOLD; http://www.boldsystems.org/) using the user web interface available after registration. The submission requires first the submission of the specimen data using the Excel-based spreadsheet following the Specimen Data Submission Protocol (http://www.boldsystems.org/index.php/resources/handbook?chapter=3_submissions.html§ion=data_submissions). To prevent the re-typing of the specimen data again, we wrote an R script converting the data from DwC to the format required for BOLD submissions (see Suppl. material
Original photo-documentation. The original photo-documentation includes the unedited high-resolution versions of photos and SEM micrographs that we used in this publication, plus many photos and SEM micrographs that were taken for comparative purposes but are not published here. We submitted all these files to Zenodo as a part of the .zip file containing all our primary data. Since the images are not easy to see in this way, we also submitted all photos to Flickr photo hosting service where they can be easily displayed; they are available at https://www.flickr.com/photos/142655814@N07/collections/72157678126129411/.
Cercyon
Leach, 1817: 95. - Type species: Dermestes melanocephalus Linnaeus (designated by
Cercyon can be distinguished from other hydrophilid genera occurring in the Greater Antilles by the following combination of characters: antenna with compact club; prothorax with conspicuous antennal groove not reaching pronotal margin; medial part of prosternum not demarcated from lateral parts; metaventrite without arcuate lines in anterolateral corners; mesoventral plate fusiform, narrowing anteriorly and posteriorly, touching anterior margin of metaventrite in one point.
Cercyon species are very similar to the members of Pelosoma, a Neotropical genus that is recorded from the Lesser Antilles; Pelosoma differs from Cercyon by the mesoventral plate widely contacting the metaventrite (it only narrowly contacts it in Cercyon). Small species of Cercyon may resemble the members of Oosternum, which can be easily distinguished from Cercyon by possessing a metaventrite with an arcuate ridge delimiting its anterolateral corner, and in some species also by elevated median part of the prosternum.
1 | Small species, body length 1.0–2.1 mm. Metaventrite with complete femoral lines (Fig. |
Cercyon nigriceps (Marsham) |
– | Larger species, body length 2.3–4.1 mm. Metaventrite without femoral lines (Figs |
2 |
2 | Mesoventral plate very wide, 1.9× as long as wide (Fig. |
Cercyon floridanus Horn |
– | Mesoventral plate moderately to very narrow, 3.3–5.9× as long as wide (Figs |
3 |
3 | Mesoventral plate wide, 3.3× as long as wide (Fig. |
Cercyon praetextatus (Say) |
– | Mesoventral plate narrow, 5.7–6.3× as long as wide as long as wide (Figs |
4 |
4 | Metaventrite with raised pentagonal area markedly wide at midlength, 0.6× as long as wide (Fig. |
Cercyon quisquilius (Linnaeus) |
– | Metaventrite with raised pentagonal area rather narrow at midlength, 0.9–1.2× as long as wide (Figs |
5 |
5 | Dorsal surface of head black, with reddish-brown spot(s) at vertex. Pronotum (Fig. |
Cercyon insularis Chevrolat |
– | Dorsal surface of head including vertex black, anterolateral margins of clypeus yellowish. Pronotum either black with yellowish to reddish lateral margins (Figs |
6 |
6 | Pronotum uniformly light brown, elytra more or less uniformly greyish-brown (Fig. |
Cercyon spiniventris sp. n. |
– | Pronotum and elytra black with pale contrasting markings (Figs |
7 |
7 | Prosternum with median ridge forming a small rounded to weakly pointed process (Fig. |
Cercyon sklodowskae sp. n. |
– | Prosternum with median ridge forming a large rounded knob (Figs |
Cercyon gimmeli species group...8 |
8 | Median lobe strongly acuminate at apex (Fig. |
Cercyon taino sp. n. |
– | Median lobe blunt at apex (Fig. |
9 |
9 | Median lobe wide basally, narrowing apically, with large gonopore and many spines in apical fifth (Fig. |
C. armatipenis sp. n. |
– | Median lobe narrowly parallel–sided, with indistinct gonopore and without spines at apex (Fig. |
C. gimmeli sp. n. |
This species group is composed of three very closely related species endemic to the island of Hispaniola. Specimens are indistinguishable on the basis of external morphology and can be only told apart by examination of male genitalia. At least in one locality, two species of this group were collected syntopically. For this reason, we refrain from using female specimens as paratypes unless the specimen was associated with males by DNA barcode, and female specimens are not listed in the text below neither in the DarwinCore spreadsheet submitted to GBIF for that reason. Below, we provide a diagnosis allowing to separate all species of the species group from other Cercyon species. Further on, we describe Cercyon gimmeli sp. n. in full, and provide only comparative diagnoses for the other two species, C. armatipenis sp. n. and C. taino sp. n.
Diagnosis of the Cercyon gimmeli species group. Members of the species group can be differentiated from other Greater-Antillean Cercyon by the following combination of characters: size 2.8–3.5 mm; dorsal surface of head black, with yellowish anterolateral margins of clypeus; pronotum black with sharply defined creamy-white areas at lateral margins; elytra black, with large, pale, rather sharply defined spot in posterior third of both elytra (Fig.
By the dorsal coloration, the species of C. gimmeli group could be confused with C. praetextatus (Say), C. floridanus Horn and C. sklodowskae sp. n. However, they can be easily distinguished from from C. praetextatus and C. floridanus Horn by the distinctly narrower mesoventral plate (compare Fig.
Distribution. The species group is endemic to Hispaniola and seems widespread on the island. No records are known from Haiti, likely due to collecting bias.
GANTC002-17 to GANTC006-17
BOLD:ADF7790.
Dominican Republic, Samaná Province, Monumento Natural Salto, El Limón 2.8 km SSW of El Limón, 19°16.56'S 69°26.47'W, 160 m a.s.l.
Holotype (male): “DOMINICAN REP.: Samaná, MN Salto El Limón 2.8 km SSW of El Limón; 19°16.56'S 69°26.47'W; 160 m; 2.ix.2014, Deler, Fikáček, Gimmel. DR29a // secondary vegetation and tiny remnants of forest among coffee plantations and pastures: in horse excrement” (
Externally identical with other members of the Cercyon gimmeli species group, it may be only distinguished from them by the morphology of the aedeagus (Fig.
Body. (Fig.
Head. Clypeus with rather sparse and shallow punctation consisting of crescent-shaped setiferous punctures intermixed with denser, smaller and rather transverse non–setiferous punctures; interstices without microsculpture. Anterior margin of clypeus with a narrow bead. Frontoclypeal suture conspicuous as a zone without punctuation, vanished in middle. Frons with punctation similar to that on clypeus, punctures of same shape all over, slightly sparser on sides; interstices without microsculpture. Eyes rather small; interocular distance about 6× the width of one eye in dorsal view. Labrum membranous, nearly completely concealed under clypeus, only with narrowly exposed sinuate anterior margin. Mentum (Fig.
Prothorax. Pronotum transverse, widest at base 2.1–2.2× wider than long; 1.8× wider at base than between anterior angles, 1.8× wider than head including eyes, as convex as elytra in lateral view. Punctation moderately dense and shallow, consisting of crescent-shaped setiferous punctures intermixed with denser, smaller and rather transverse non–setiferous punctures; punctures slightly feebler on sides. Prosternum (Fig.
Pterothorax. Scutellar shield 1.1× as long as wide, sparsely punctured. Elytra widest at anterior fifth, 2.55–2.85× as long as pronotum, 1.25–1.35× as wide as pronotum; surface glabrous (Fig.
Legs. Femora with sparse rather shallow punctures ventrally, interstices with weak microsculpture at bases, consisting of longitudinal lines; tibial grooves distinct. Tibiae with rather large lateral spines. Metatibiae moderately narrow and elongate, slightly bent outwards, 0.3–0.4× as long as elytra, 5.3× as long as wide. Metatarsus long, 0.9× as long as metatiba, with few short but rather stout setae ventrally.
Abdomen with five ventrites, first abdominal ventrite about as long as second and third ventrites combined, with distinct median longitudinal carina (Fig.
Male genitalia. Median projection of sternite 9 (Fig.
In some specimens the pale spots at sides of the scutellar shield are longer, almost reaching the second fourth of elytral length.
We are pleased to dedicate this species to Matthew L. Gimmel (Santa Barbara Museum of Natural History), who participated in the expedition to the Dominican Republic and collected part of the type series of this species.
Dominican Republic: Barahona, La Vega, Monseñor Nouel, Samaná (Fig.
All specimens were collected in in broad-leaf tropical forests and coffee plantations and pastures, mainly on cow and horse dung, but also on human dung, rat carrion, at black-light or at day by beating.
GANTC013-17, GANTC014-17
BOLD:ADF5573
Dominican Republic, Independencia Province, Parque Nacional Sierra de Neiba, 11.3 km NW of La Descubierta, 18°39.81'N, 71°46.17'W, 1650 m a.s.l.
Holotype (male): “DOMINICAN R.: Independencia, PN Sierra de Neiba, 11.3 km NW of La Descubierta; 1650 m, 18°39.81'N, 71°46.17'W; 18.viii.2014, Deler, Gimmel DR13 // disturbed montane cloud forest with many ferns and mosses: in cow excrement” (
Externally identical with other members of the Cercyon gimmeli species group, it may be only distinguished from them by the morphology of the aedeagus (Fig.
Measurements. (Fig.
Conforming to the description of C. gimmeli, with the following differences: Pterothorax. Punctation of even intervals consisting of small non-setiferous punctures, but here and there with infrequent crescent-shaped larger setiferous punctures; only lateralmost elytral interval completely pale.
Male genitalia. Median projection of sternite 9 (Fig.
The species name is derived from Latin words armatus (armed) and penis (penis), in reference to the diagnostic character of this species, i.e. the apex of the median lobe armed by small spines.
Dominican Republic: Independencia (Fig.
Specimens were collected in cow dung in a cloud forest.
GANTC007-17 to GANTC009-17
BOLD:ADF5574
Dominican Republic, Samaná Province, dam 2.5 km N of Samaná, 58 m a.s.l., 19°13.70'N, 69°19.85'W.
Holotype (male): “DOMINICAN REP.: Samaná, dam 2.5 km N of Samaná, 19°13.70'N, 69°19.85'W; 58 m, 5.ix.2014, Deler, Fikáček, Gimmel DR35 // in older cow excrements dampered by recent rains at the grassy bank of a reservoir” (
Externally identical with other members of the Cercyon gimmeli species group, it may be only distinguished from them by the morphology of the aedeagus (Fig.
Measurements. (Fig.
Conforming with description of C. gimmeli, with the following differences: Pterothorax. Punctation of even intervals consisting of small non-setiferous punctures, but here and there with infrequent crescent-shaped larger setiferous punctures; only lateralmost elytral interval completely pale.
Male genitalia. Median projection of sternite 9 (Fig.
The new species is named after the indigenous Taíno people inhabiting the Greater Antilles including Hispaniola before and at the time of the arrival of Europeans.
Dominican Republic (Samaná, La Vega, Independencia) (Fig.
Examined specimens and the associated larvae (see below) were collected from leaf litter of montane forests and from cow dung.
Holotype (male): “JAM., St. Thomas, Corn Puu Gap [sic!, = Corn Puss Gap], 2100’, 4mi. N, Bath, 3-8.VIII.1974, S. Peck, dung” (
Cercyon sklodowskae sp. n. can be easily differentiated from other Greater-Antilles Cercyon species by the following combination of characters: body size 3.1–3.5 mm; dorsal surface of head black with yellowish anterolateral margins of clypeus; pronotum black with sharply defined yellowish areas at lateral margins (Fig.
C. sklodowskae may be confused with C. praetextatus (Say) and Cercyon floridanus Horn and the members of C. gimmeli species group. It can be distinguished from C. praetextatus and C. floridanus by the head with widely yellowish clypeal margin and yellow coloration of elytra expanded basally (clypeus black and basal part of elytra only narrowly yellow in C. praetextatus and C. floridanus) and by much narrower mesoventral plate (5.8× as long as wide in C. sklodowskae, 1.9× as long as wide in C. floridanus and 3.3× in C. praetextatus). It differs from members of C. gimmeli group by the mesoventral plate reaching anterior part of metaventrite (not reaching anterior margin in C. gimmeli group), only a small rounded process of median prosternal ridge (large rounded knob in C. gimmeli group), straight metatibia (slightly curved outside in C. gimmeli group), and last abdominal ventrite in females forming a triangular projection (without projection in C. gimmeli group). It differs from all these species by the morphology of male genitalia; by the apically spinose median lobe it resembles C. armatipenis, but differs from it by parameres much shorter than phallobase.
Body. (Fig.
Head. Clypeus with moderately dense and shallow punctation consisting of crescent-shaped punctures intermixed with denser, slightly smaller and rather transverse punctures; interstices without microsculpture. Anterior margin of clypeus with a narrow bead. Frontoclypeal suture conspicuous as a zone without punctation, vanishing mesally. Frons with punctation similar to that on clypeus, punctures sparser on sides; interstices without microsculpture. Eyes rather small; interocular distance about 6× the width of one eye in dorsal view. Labrum membranous, nearly completely concealed under clypeus, only with narrowly exposed sinuate anterior margin. Mentum (Fig.
Prothorax. Pronotum transverse, widest at base 2.1–2.2× wider than long; 1.6–1.7× wider at base than between front angles, 1.7× wider than head including eyes, as convex as the elytra in lateral view. Punctation (Fig.
Pterothorax. Scutellar shield 1.25× as long as wide, sparsely punctured. Elytra widest at anterior fifth, 2.7–3.0× as long as pronotum, 1.2× as wide as pronotum, surface (Fig.
Legs. Femora with sparse rather shallow punctures ventrally, interstices with weak microsculpture consisting of longitudinal lines; tibial grooves distinct. Tibiae with rather small lateral spines. Metatibiae moderately narrow and elongate, straight, 0.3–0.4× as long as elytra, 5.3× as long as wide. Metatarsus long, 0.9× as long as metatibia, with short rather stout setae ventrally.
Abdomen. With five ventrites, first abdominal ventrite (Fig.
Genitalia. Sternite 9 (Fig.
We dedicate this species to the eminent physicist and chemist Marie Skłodowska-Curie, on whose honor the Marie Skłodowska-Curie actions program of the European Union, funding this research, is named.
Jamaica: Saint Thomas (Fig.
The specimens were collected on dung and using dung and carrion-baited traps in cloud forests.
Cercyon floridanus Horn, 1890: 303.
Cercyon
floridanuus
Smetana (
Florida (without specific locality).
CAYMAN ISLANDS: Grand Cayman: blacklight trap, vi.1992, leg. blacklight trap (1 male;
GRAND CAYMAN: 3 km W Colliers, 19°21'N, 81°07'W (
Body size 2.35–2.70 mm; dorsal surface of head (Fig.
Cercyon floridanus is part of the C. tristis group according to
Cercyon floridanus is distributed in the southeastern USA, mainly in Florida, but rare records are also known from Georgia, Louisiana and Mississippi (
Sphaeridium praetextatum Say, 1825: 190.
Cercyon praetextatum (Say): Melsheimer, 1853: 37.
For complete synonymy see
USA, “Cambridge” (based on neotype designated by
Greater Antillean specimens studied. CUBA: Santiago de Cuba: Dos Caminos, farm field, MV lights, 20.18043°N, 75.77806°W, 165 m, 23.iii.2013, leg. A. Smith & A. Deler-Hernández (1 spec.:
CUBA: Habana Province: Laguna de Ariguanabo (
Body size 2.7–4.1 mm; dorsal surface of head (Fig.
This species was assigned to the C. marinus group according to
Cercyon praetextatus is widely distributed in North America (southern Canada, USA, Mexico;
This species seems to prefer wet environments, living primarily on many kind of organic debris, like decomposing plant remnants, carrion and dung (
GANTC010-17
BOLD:ADF5572
Figures in Flickr: www.flickr.com/photos/142655814@N07/albums/72157671689463811
Dominican Republic, Monseñor Nouel Province, Parque Nacional La Humeadora; 11.6 km SSW of Piedra Blanca, 636 m a.s.l., 18°44.92'N, 70°21.63'W.
Holotype (male): “DOMINICAN REP.: Msñ. Nouel, PN La Humeadora; 11.6km SSW, of Piedra Blanca; 18°44.92'N, 70°21.63'W; 636 m; 8.ix.2014, Deler, Fikáček, Gimmel DR41 // in horse excrement in moist broad-leaf forest in a valley of a small stony stream” (
Body size 3.4–4.1 mm; dorsal surface of head black with yellowish anterolateral margins of clypeus (Fig.
Cercyon spiniventris somewhat resembles C. nigriceps by the dorsal coloration pattern (predominantly black head and rather homogeneously brown pronotum and elytra); it can be easily distinguished from C. nigriceps by much larger body size (3.4–4.1 mm in C. spiniventris, 1.0–2.1 mm in C. nigriceps) and by the lack of femoral lines on the metaventrite (present in C. nigriceps). Cercyon spiniventris is unique among Caribbean Cercyon species by a presence of a long spiniform process in the first abdominal ventrite of females.
Body (Fig.
Head. Clypeus with dense and moderately deep punctation consisting of crescent-shaped setiferous punctures intermixed with denser, smaller and rather transverse non-setiferous punctures; interstices without microsculpture. Anterior margin of clypeus with a narrow bead. Frontoclypeal suture conspicuous as a zone without punctuation, vanished mesally. Frons with punctation similar to that on clypeus, punctures of same shape all over; interstices without microsculpture. Eyes rather small; interocular distance about 5.4× the width of one eye in dorsal view. Labrum membranous, nearly completely concealed under clypeus, only with narrowly exposed sinuate anterior margin. Mentum (Fig.
Prothorax. Pronotum transverse, widest at base 2.1–2.3× wider than long; 1.7× wider at base than between anterior angles, 1.8× wider than head including eyes, as convex as elytra in lateral view. Punctation rather dense and moderately deep, consisting of crescent-shaped setiferous punctures intermixed with denser, smaller and rather transverse non-setiferous punctures; punctures slightly feebler on sides. Prosternum (Fig.
Pterothorax. Scutellar shield about as long as wide, moderately densely punctured. Elytra widest at anterior fifth, 2.7–2.9× as long as pronotum, 1.1–1.2× as wide as pronotum, surface (Fig.
Legs. Femora with sparse rather shallow punctures ventrally, interstices with weak granulose microsculpture; tibial grooves distinct. Tibiae with moderately large lateral spines. Metatibiae moderately narrow and elongate, slightly bent outwards, 0.4× as long as elytra, 6.0× as long as wide. Metatarsus moderately long, 0.7–0.8× as long as metatibia, with short rather stout setae ventrally.
Abdomen with five ventrites, first abdominal ventrite longer than second and third ventrites combined, with long setae in medial third, median longitudinal carina present, slightly narrowing posteriad, not projecting posteriorly in males, projecting posteriad as a short spine in females (Fig.
Genitalia. Median projection of sternite 9 (Fig.
The name of this species is derived from Latin words spina (spine) and venter (underside), in reference to the spine-like process on the first abdominal ventrite of females.
Dominican Republic: Duarte, Independencia, La Vega, Monseñor Nouel, Samaná (Fig.
Most of the specimens were collected in cow and horse dung in tropical forest and surrounding pastures.
Dermestes nigriceps Marsham, 1802: 72.
Cercyon nigriceps Stephens (1829: 151).
= Dermestes atricapillus Marsham, 1802: 72 (synonymized by Gemminger and Harold 1868: 498; precedence of C. nigriceps over C. atricapillus determined by Stephens 1939: 97, see also
= Sphaeridium centrimaculatum Sturm, 1807: 23 (synonymized by Gemminger and Harold 1868: 498).
= Cercyon striatus Sharp, 1882: 108 (synonymized by
= Cercyon panamensis Hansen, 1999: 286 (replacement name of C. striatus Sharp; synonymized by
For complete synonymy see
GANTC015-17
BOLD:AAO0116
“Britannia” [= Great Britain, without specified locality].
CAYMAN ISLANDS: Cayman Brac: black-light trap, 06.vi.2008, lgt. R.H. Turnbow & B.K. Dozier (3 spec.:
CUBA: Matanzas: Cárdenas (as C. centrimaculatum,
Body size 1.0–2.1 mm; dorsal surface of head black; pronotum (Fig.
This species was assigned to C. nigriceps group (= C. atricapillus group) by
This is an adventive species currently distributed in all zoogeographical regions. In Greater Antilles widespread in all islands: Cayman Islands, Cuba (Artemisa, Cienfuegos, Guantánamo, Holguín, Matanzas, Santiago de Cuba;
A terrestrial species collected in cow and horse dung and in decaying plant matter (e.g., compost piles). It is also frequently collected at light.
Scarabaeus quisquilius Linnaeus, 1761: 138.
Cercyon quisquilium Stephens (1829: 153).
For complete synonymy see
“Suecia” [= Sweden, without specified locality].
CUBA: Holguín: Mayarí Municipality, Feltón, Vuelta Larga, permanent lagoon, 23.iii.2013, leg. A. Deler-Hernández (2 spec.:
JAMAICA: without precise locality (
Body size 2.4–3.2 mm; dorsal surface of head completely black; pronotum (Figs
Cercyon quisquilius was assigned to C. unipunctatus group according to
Cercyon quisquilius is a species native to the Palearctic Region, but currently introduced to the Nearctic, Neotropical and Australian Regions (
Cercyon insulare Chevrolat, 1863: 208.
Cercyon
insulare
GANTC001-16, GANTC011-17, GANTC012-17
BOLD:ADC9388.
Cuba: Havana.
Holotype (unsexed specimen): “Havana, D. Poey // Cercyon insulare, Chev Cuba, [illegible] // TYPE [red label]” (
Cercyon insularis Chevrolat and C. variegatus Sharp. a–i C. insularis: a–c dorsal, ventral and lateral habitus of the non-type specimen from Cuba d habitus of the holotype e labels of the holotype f–i male genitalia of non-type specimen from Dominican Republic. j–o lectotype of C. variegatus: j dorsal habitus k labels l–o male genitalia. Genital parts illustrated: f, l tegmen of aedeagus g, m median lobe of aedeagus h, n detail of apex of median lobe; i, o 9th sternite.
CUBA: Camagüey: Sierra de Cubitas municipality, Limones-Tuabaquey, 21°35'52.10"N, 77°47'17.62"W, 16.v.2013, leg. R. Anderson (1 spec.:
(as C. variegatus). JAMAICA: without precise locality (
Body size 2.4–3.4 mm; dorsal surface of head (Fig.
Body. (Fig.
Head. Clypeus with moderately dense and shallow punctation consisting of small transverse punctures; interstices without microsculpture. Anterior margin of clypeus with narrow bead. Frontoclypeal suture conspicuous as a zone without punctuation, vanished mesally. Frons with punctation similar to that on clypeus, punctures sparser on sides; interstices without microsculpture. Eyes rather small, interocular distance about 6× the width of one eye in dorsal view. Labrum membranous, nearly completely concealed under clypeus, only with narrowly exposed sinuate anterior margin. Mentum (Fig.
Prothorax. Pronotum transverse, widest at base 2.1–2.2× wider than long; 1.7–1.8× wider at base than between front angles, 1.8× wider than head including eyes, as convex as pronotum in lateral view. Punctation rather dense and moderately deep, consisting of crescent-shaped punctures intermixed with denser, slightly smaller and rather transverse punctures; punctures slightly feebler on sides. Prosternum (Fig.
Pterothorax. Scutellar shield 1.2× as long as wide, sparsely punctured. Elytra widest at anterior fifth, 1.0–1.1× longer than wide, 2.6–2.8× as long as pronotum, 1.2–1.3× as wide as pronotum, surface glabrous, with 10 series of punctures; series 6, 8 and 9 not reaching anterior margin, surface glabrous (Fig.
Legs. Femora with sparse shallow punctures ventrally, interstices with weak microsculpture consisting of longitudinal lines; tibial grooves distinct. Tibiae with rather small lateral spines. Metatibiae moderately broad and long, straight, 0.33× as long as elytra, 5× as long as wide. Metatarsus long, 0.86–0.89× as long as metatibia, with just a few short rather stout setae ventrally.
Abdomen with five ventrites, first abdominal ventrite (Fig.
Male genitalia. Median projection of sternite 9 (Fig.
The general dorsal coloration of the pronotum and elytra varies from yellow to dark reddish-brown. In dark specimens, lateral pronotal spots join the large central spot, and the whole anterior part of elytra and the elytral suture are distinctly darkened, with pale areas maintained in humeral area and at sides of scutellar shield. In pale specimens, the lateral pronotal spots are rather small and sometimes very vague and indistinct, and the elytra are completely yellow except base, sutural interval and the posthumeral dark spots.
Cercyon insularis seems to be widely distributed across Greater and Lesser Antilles, here we are recording it from Cuba, Dominican Republic, Puerto Rico, Grenada, Saint Lucia and Dominica. It seems that all records of C. variegatus from the Caribbean (Jamaica, Puerto Rico:
Most of the specimens were collected in cow and horse dung on pastures, in coffee plantations and in tropical forests; few were collected using flight intercept traps.
Cercyon insularis was only briefly mentioned once by
Cercyon insularis and C. variegatus belong to a species complex corresponding to the C. variegatus group of
DOMINICAN REPUBLIC: La Vega: PN Valle Nuevo, Salto Aguas Blancas, sifting of moist leaf litter in small remnants of montane forest in a small ravine with a spring and on slopes just above the small river, 18°50.60'N, 70°40.68'W, 1655 m a.s.l., 25.viii.2014, leg. Deler, Fikáček & Gimmel (DR21) (2 larvae associated with adults:
Head capsule (Fig.
PUERTO RICO: Naguabo: El Yunque National Forest, (southern part), 3.45 km N of Río Blanco at road PR191, in horse excrements on exposed small pasture on the slope of El Yunque massive, 18°14.8'N, 65°47.9'W, 170 m a.s.l., 24.vi.2016, leg. Deler-Hernández, Fikáček & Seidel (PR2a) (2 larvae associated with adults:
Head capsule (Fig.
Partial COI sequences of the 19 Cercyon specimens sampled resulted in a 610 bp alignment. JModelTest (
Faunal composition of Cercyon in the Greater Antilles. Our revision raises the number of Cercyon species from six to 10, and shows that the composition of the fauna largely differs from the original ideas: (1) five species, i.e. half of the fauna, are probably single-island endemics (four species in Hispaniola, one in Jamaica) and one additional species (C. insularis) seems to be a widespread Caribbean endemic not occurring in the mainland Americas; (2) only two species occurring in the Greater Antilles (C. praetextatus and C. floridanus) are native to the American continent, of which C. floridanus is moreover limited only to Cayman islands and does not occur in the four main islands of Greater Antilles; (3) two introduced Old World species (C. nigriceps and C. quisquilius) occur in the Greater Antilles, both of which are nowadays also widespread in the American continent (
The comparison of the original ideas about the Cercyon fauna and its real composition corresponds to the situation found in many other groups of minute arthropods, which were recently studied in the Greater Antilles. These studies frequently discover higher numbers of single-island endemics than expected, and reveal that some widespread continental species recorded from the Caribbean are in fact endemic species closely related but not identical to the continental ones (e.g.,
The probable single-island endemics discovered during this study are distributed in Hispaniola (four species) and Jamaica (one species) only. No new species were discovered in Cuba and Puerto Rico, despite our collecting effort in both islands. Our field work in Puerto Rico was rather short and did not cover all mountain regions. Therefore, we cannot exclude additional discoveries in the island. On the other hand, our sampling effort was highest in Cuba, with intensive field work performed in 2010–2016. Hence we consider the discovery of a new endemic species in Cuba less likely. This puts in contrast Cuba, i.e. the largest Greater Antillean island, with no endemic species, with the smaller Hispaniola, hosting at least four single-island endemics. Moreover, the discovery of C. armatipenis sp. n. in a single locality in Sierra de Neiba at the Dominican-Haitian border indicates that additional species may be expected in this species complex in the western part of the island (i.e. Haiti) which was not sampled so far. In the same manner, there is a chance that the only species so far known exclusively from the Lesser Antilles, Cercyon cribratus
Both the extremely similar morphology and results of the analysis of the COI sequences imply that the C. gimmeli species complex forms a monophyletic clade endemic to Hispaniola. The fact that all species of the complex occur sympatrically and no clear geographic pattern can be observed in their ranges suggests that the radiation of this group in Hispaniola may be a result of subdivision of Hispaniola in smaller paleoislands during the Oligocene to Middle Miocene followed by range expansion when the paleoislands got interconnected more recently (
Novel morphological characters. Morphological studies of the Greater-Antillean Cercyon revealed some characters of adults relevant to species discrimination and identification, but not used before: the presence/absence and shape of the projection of the anterior part of the prosternal medial ridge, and the sexually dimorphic characters of abdominal ventrites found in C. spiniventris and C. sklodowskae. Females of C. spiniventris are characterized by a spine-like process on the first abdominal ventrite (absent in males), which has not been recorded in any other Cercyon species so far, but is present in females of the Australian megasternine genus Cercyodes Broun (
Interesting discoveries were also made by a brief examination of the larvae of C. insularis and C. taino. Both larvae are surprisingly quite different from each other, differing especially in the head proportions, extent of leg reduction, shape of the tergite of 8th abdominal segment, and surface vestiture membranous parts of thorax and abdomen. In all these characters, C. insularis is more similar to other Cercyon larvae described in the literature (C. quisquilius and C. praetextatus:
The novel characters mentioned above of both adults and larvae are useful for diagnostic purposes, but can also have phylogenetic signal which will help to corroborate the results of the ongoing phylogenetic study of the tribe Megasternini. The differences found between the larvae of C. insularis and C. taino and the differences in the number of lenses between different megasternine taxa show that larval morphology of the Megasternini is not that uniform as previously believed, and is in need of more studies.
Subgeneric assignment of the Greater Antilles Cercyon species. All species treated in this paper fall into the concept of Cercyon sensu stricto. However, we refrain from assigning them to any subgenus since the systematics of the genus Cercyon and allies is currently under study, and previous studies have shown that Cercyon in the current concept may be a polyphyletic assemblage of species (e.g.
ADH, MF and MS performed the field work; EAV and MF accumulated additional museum material; EAV performed the majority of morphological studies, prepared the first draft and photodocumentation; EAV and MS did the molecular work, analysis of the data and prepared the data for submissions to BOLD; VS, MF and EAV prepared the datasets, wrote and tested the scripts, and submitted the data to BOLD, GBIF and Flickr; all authors commented drafts of the paper at different stages and helped with completing the manuscript for submission.
We greatly appreciate the help of all the curators who allowed us to work on the specimens held in their collections. Matthew L. Gimmel (Santa Barbara Museum of Natural History) in sincerely ackowledged for their contribution during the fieldwork performed in the Dominican Republic. This work was supported by the European Union's Horizon 2020 research and innovation program under the Marie Skłodowska-Curie grant agreement No. 642241 to M. Seidel and E. Arriaga-Varela, and the Ministry of Culture of the Czech Republic (DKRVO 2017/14, National Museum, 00023272) to Martin Fikáček. The work of the first three authors at the Department of Zoology, Charles University in Prague was partly supported by grant SVV 260 434 /2017. The Synthesys project. http://www.synthesys.info/ funded the stay of EA-V at the
Darwin Core formatted distribution data
Data type: species distribution
Explanation note: The Excel file includes all fields (columns) defined by the DarwinCore (DwC) format relevant to our data (headers highlighted in green) and ordered in a customized way (order of columns is not important, columns are identified by their header names that need to remain unchanged). We added a few fields (columns), which are not present in DwC, to include additional information needed in our case (about specimens in a DNA-tissue collection, DNA extracts done, and DNA barcodes), or to make the data input more convenient (two separate columns for starting and ending event dates, from which the date range in DwC format is formed and saved in the eventDate field); headers of all additional fields are marked in grey in Suppl. material
Genetic distances within and between species of Caribbean Cercyon
Data type: molecular data
Explanation note: Mean genetic distances based on barcode CO1 sequences within (in grey fields) and between species (in white fields) calculated using the Maximum Composite Likelihood model as implemented in MEGA7. Standard error estimates (in square brackets) obtained by bootstraping (1000 replicates).@.
DwC2BOLD script
Data type: script
Explanation note: We used this script to automatically transform the DwC-formatted specimen data into the format required for the submission of specimen data to BOLD. The script uses the Barcode field (column) in our DwC spreadsheet to filter out the specimens that were barcoded and that need to be submitted to BOLD (notice that the Barcode field was added by us and is not a part of the DwC format!). Due to the slightly different data format used in DwC and BOLD, it is not possible to convert all information automatically, and the resulting BOLD-formatted data should be checked before submission. The script displays alerts for fields in which a mismatch between DwC and BOLD is possible. The script is available in Suppl. material
DwC2Map script
Data type: script
Explanation note: The script reads the decimal GPS data from our DwC Excel file and creates a dataframe with all observations for each species. In the next step it allows to construct the maps of the area from freely available on-line data and map the occurrences of thes species on these maps. The maps in Figs