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A Chromosomal Analysis of 21 Species of Oniticellini and Onthophagini (Coleoptera: Scarabaeidae)

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C. J. WILSON & R. B. ANGUS School of Biological Sciences, Royal Holloway, University of London, UK A CHROMOSOMAL ANALYSIS OF 21 SPECIES OF ONITICELLINI AND ONTHOPHAGINI (COLEOPTERA: SCARABAEIDAE) Wilson, C. J. & R. B. Angus, 2005. A chromosomal analysis of 21 species of Oniticellini and Onthophagini (Coleoptera: Scarabaeidae). – Tijdschrift voor Entomologie 148: 63-76, figs. 1-6, tables 1-2. [ 0040-7496]. Published 1 June 2005. The karyotypes of 2 species of Oniticellini and 19 species of Onthophagini are described and illustrated. Euonthophagus pallipes, Caccobius schreberi, Euonthophagus amyntas, E. atramentarius, Onthophagus taurus, O. hirtus, O. furcatus, O. stylocerus, O. vacca, O. similis, O. opacicollis, O. coenobita, O. lucidus, O. ovatus, O. joannae, O. ruficapillus, O. gazella and O. albicornis all have karyotypes with 2N = 18 + Xy. Only female karyotypes have been obtained for Euoniticellus fulvus, Onthophagus illyricus and O. maki. These species have 2N = 20 chromosomes, including the unidentified sex chromosomes. B-chromosomes have been found in Euonthophagus amyntas, Onthophagus vacca, O. similis and O. gazella, and chromosome polymorphisms result- ing from pericentric inversions have been found in Onthophagus vacca and O. albicornis. Centromeric C-bands, of varying sizes, are present in nearly all the chromosomes, and the species of Euoniticellus and Euonthophagus have some chromosomes with a totally heterochro- matic arm, a feature not encountered in the Onthophagus species. The three species of the Onthophagus ovatus group have karyotypes clearly distinct from one another, as do O. similis and O. opacicollis. However, no difference has been found between the karyotypes of O. taurus and O. illyricus. C. J. Wilson & R. B. Angus, School of Biological Sciences, Royal Holloway, University of London, Egham, Surrey TW20 0EX, UK. E-mail: [email protected] Key words. – Chromosomes; karyotypes; C-banding; B-chromosomes; Euoniticellus; Caccobius; Onthophagus; Scarabaeidae. In the course of research for a Ph.D. degree of origin is given in table 1. British localities are (Wilson 2002) chromosomal data on 18 species referred to their Watsonian Vice-Counties, as set out of Euoniticellus, Caccobius, Euonthophagus and by Dandy (1969). French localities are listed by Onthophagus were accumulated, and continuing in- Département, written in full. Spanish localities are vestigations by R. B. Angus have since added a listed by Provincia, written in full, as are provinces in further three Onthophagus species to the list. There is Cyprus (Eparkhia) and the Dominican Republic at present no published information on the chromo- (Provincias). The number of specimens from which somes of ten of the species, and where information is successful preparations were obtained is given in available it very seldom includes karyotypes, the usu- parentheses after the list of localities for each species. al information being the number of chromosomes The methods of chromosome preparation and cura- and/or the meioformula (the arrangement of the tion of the material are given by Wilson (2001). chromosomes at first metaphase of meiosis). The Relative Chromosome Length () is the length of material presented here therefore provides a first each chromosome expressed as a percentage of the to- opportunity for a comparison of the karyotypes of a tal haploid autosome length in the nucleus. It is used, suite of small scarabaeid dung beetles. without statistical analysis, to give an account of the approximate sequence of chromosome lengths for the different species. Where only females are available the M sex chromosomes cannot be identified, so total A list of beetle species from which chromosome haploid autosome length is not available. In these preparations have been obtained, and their localities cases a ‘standard length’ () is used, using the total 63 Downloaded from Brill.com09/30/2021 07:32:02AM via free access T E, 148, 2005 Table 1. Material studied and taxonomy. Scientific name Localities (number of specimens analysed) Oniticellini Euoniticellus Janssens, 1953 E. fulvus (Goeze, 1777) .......................... Indre-et-Loire: Souvigné; Indre: La Brenne. (3 specimens.) E. pallipes (Fabricius, 1881) .................... Limassol: Akrotiri, Fassouri. (3 specimens.) Onthophagini Caccobius Thomson, 1859 C. schreberi (Linnaeus, 1758) .................. Cáceres: Abadía. (2 specimens.) Euonthophagus Balthasar, 1959 E. amyntas (Olivier, 1789) ...................... Cáceres: Abadía. (1 specimen.) E. atramentarius (Ménétriès, 1832) ........ Pafos: Anadhiou; Agia Varvara. (7 specimens.) Onthophagus Latreille, 1802 subgenus Onthophagus s. str. O. taurus (Schreber, 1759) ..................... Indre: La Brenne. Málaga: Torrox. (5 specimens.) O. illyricus (Scopoli, 1763) ..................... Limassol: Akrotiri, Fassouri. (3 specimens.) subgenus Trichonthophagus Zunino, 1979 O. hirtus (Illiger, 1803) .......................... Cádiz: Tarifa. ( specimens.) O. maki (Illiger, 1803) ........................... Cádiz: La Linea. (1 specimen.) subgenus Furconthophagus Zunino, 1979 O. furcatus (Fabricius, 1781) .................. Cádiz: Tarifa. (1 specimen.) Subgenus Palaeonthophagus Zunino, 1979 O. stylocerus Graëlls, 1851 ...................... León: Sanabria. Cantabria: Corconte. (2 specimens.) O. vacca (Linnaeus, 1767) ...................... East Sussex: Rye; Middlesex: Staines Moor. Indre: La Brenne; Maine-et-Loire: Le Lion d’Angers. Cantabria: Corconte. (8 specimens.) O. similis (Scriba, 1790) ......................... North Devon: Ilfracombe; East Sussex: Rye; Surrey: Ockham Common; Worcestershire: Wyre Forest. (8 specimens.) O. opacicollis Reitter, 1893 ..................... Pafos: Pafos; Kholetria; Agia Varvara; Anadhiou; Cape Drepanon. (10 specimens.) O. coenobita (Herbst, 1783) ................... East Kent: Betteshanger; Berkshire: Old Windsor; Windsor Deer Park; Worcestershire: Wyre Forest. (3 specimens.) O. lucidus (Sturm, 1800) ........................ Pafos: Anadhiou. (3 specimens). The O. ovatus group O. ovatus (Linnaeus, 1767) ..................... Indre: La Brenne; Loir-et-Cher: Pinail. (8 specimens). O. joannae Goljan, 1953 ........................ East Kent: Lydden; Surrey: Box Hill. (7 specimens.) O. ruficapillus Brullé, 1832 ..................... Limassol: Akrotiri, Fassouri. (4 specimens.) subgenus Digitonthophagus Balthasar, 1959 O. gazella (Fabricius, 1787) .................... . Espaillat: Las Flores near Santa Maria; Maria Trinidad Sanchez: Genebra near the Rio Boba; Samana: El Limón. (8 specimens.) not attributed to a subgenus O. albicornis (Beauvois, 1805) ................ . Espaillat: Las Flores near Santa Maria. (5 specimens.) haploid chromosome (not autosome) length in the dard terms denoting centromere position. Based on nucleus. In practice, the figures obtained would be Sumner (2003) these may be defined as follows: only slightly different from values. Centromere metacentric – 46 – 50; submetacentric – 26 – Index (), the length of the short arm as a percentage 45; subacrocentric – 16 – 25; acrocentric – 3 – of the total length of the chromosome is used as a 15. Unless otherwise stated in the figure captions, all measure of the location of the centromere. In most the karyotypes illustrated were obtained from mid- cases the calculated s are not given, but instead stan- gut cells. 64 Downloaded from Brill.com09/30/2021 07:32:02AM via free access W A: Chromosomes Oniticellini & Onthophagini 123456789XyB a b c d e f g h 5 µm Fig. 1. Mitotic chromosomes of Euoniticellus, Caccobius and Euonthophagus, arranged as karyotypes.– a, Euoniticellus fulvus, &, La Brenne; b, E. fulvus, &, La Brenne, C-banded; c, E. pallipes, (, Fassouri; d, Caccobius schreberi, (, testis, Abadía; e, f, Euonthophagus amyntas, (, Abadía, e plain, with one B-chromosome, f with some spontaneous C-banding and with one replicate of autosomes 4 and 5 missing; g, E. atramentarius, (, Anadhiou, with one replicate of autosome 5 missing; h, E. atramentarius, (, Anadhiou, with one replicate of autosomes 6 and 9 missing. 65 Downloaded from Brill.com09/30/2021 07:32:02AM via free access T E, 148, 2005 123456789Xy a b 10 c d Xy e f g 10 h Xy i 5 µm Fig. 2. Mitotic chromosomes of Onthophagus s. str., O. (Trichonthophagus) and O. (Furconthophagus), arranged as karyotypes. – a, Onthophagus taurus, (, Torrox; b, O. taurus, &, La Brenne; c, d, O. illyricus, &, Fassouri; e, O. hirtus, (, Tarifa; f, g, O. hirtus, &, Tarifa, e plain, f the same nucleus C-banded. One replicate of chromosome 9 is missing; h, O. maki, &, La Linea; i, O. furcatus, (, Tarifa. 66 Downloaded from Brill.com09/30/2021 07:32:02AM via free access W A: Chromosomes Oniticellini & Onthophagini R throughout their lengths. However, partial kary- otypes from mid-gut cells showed no arms with the Oniticellini chromatids close together, and no C-banded prepara- tion is available. There do not appear to be any chro- Euoniticellus fulvus mosomes with heterochromatic arms in this species. (figs. 1a, b) First metaphase of meiosis (not shown) shows the sex bivalent as a normal Xyp. Published information: 10 bivalents (() (Virkki 1954). 2N = 20 (&). The only successful preparations Euonthophagus amyntas were from females. The standard lengths of the chro- (figs. 1e, f) mosomes range from about 16 to 5. Chromosomes 1, 3 and 5 – 7 are more or less metacentric, 2 and 8 Published information: 2N = 20 (() (Virkki 1951). are subacrocentric, and the rest are more or less acro- 2N = 18 + Xy + B-chromosomes. The s of the
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  • (Coleoptera: Scarabaeoidea) Genitalia: Some Remarks to a Recent Paper

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    ISSN 0065-1737 Acta Zoológica MexicanaActa Zool. (n.s.), Mex. 30(2): (n.s.) 439-443 30(2) (2014) Nota Científica (Short Communication) ABOUT DUNG BEETLES (COLEOPTERA: SCARABAEOIDEA) GENITALIA: SOME REMARKS TO A RECENT PAPER Zunino, M. 2014. Acerca de los genitalia en escarabajos (Coleoptera: Scarabaeoidea): algunas observaciones a un artículo reciente. Acta Zoológica Mexicana (n. s.), 30(2): 439-443. RESUMEN. Se analiza críticamente una publicación extensa sobre los genitalia en diferentes grupos de Coleoptera Scarabaeinae; se subraya la necesidad de revisar las homologías de las diferentes estructuras así como de unificar la nomenclatura en una forma no arbitraria como una precondicionante para su uso en taxonomía y análisis filogenético. Male, and less frequently female, genitalia characters have often been utilized in beetles taxonomic and phylogenetic research, since as early as the beginning of the 20th century, starting at least from the seminal books by Sharp and Muir (1912), and subsequently by Jeannel (1955), where the genital anatomy of Coleoptera was first systematized (for some historical remarks mostly concerning Scarab beetles see Zu- nino 2012). The prevailing importance of genitalia not just in alpha taxonomy, but as reliable phylogenetic tracers against external morphological characters underlies a great amount of papers throughout at least fifty years; it was explicitly stressed by Zunino (1983, 1987), Zunino & Palestrini (1988), and was formally demonstrated by Tarasov & Solodovnikov (2011) by means of a cladistic analysis of a large sample of Onthophagini. Medina et al. (2013) emphasize the importance of recognizing the ho- mologies within different male genital structures in the frame of Scarabaeinae, and of unifying the relative nomenclature, for several scientific purposes.
  • The Lamellicorn Beetles of Southern Sardinia ( Coleoptera: Scarabaeoidea) *

    The Lamellicorn Beetles of Southern Sardinia ( Coleoptera: Scarabaeoidea) *

    Conservazione habitat invertebrati 5: 353–387 (2011) Cnbfvr The lamellicorn beetles of southern Sardinia ( Coleoptera: Scarabaeoidea)* Giuseppe M. CARPANETO1, Emanuele PIATTELLA2, Giovanni DELLACASA3, Marco DELLACASA4, Riccardo PITTINO5, Adriano MAZZIOTTA6 1Dipartimento di Biologia Ambientale, Università degli Studi Roma Tre, Viale Guglielmo Marconi 446, I-00146 Rome, Italy. E-mail: [email protected] 2Dipartimento di Biologia e Biotecnologie "Charles Darwin" (Entomologia), Sapienza Università di Roma, Piazzale Valerio Massimo 6, I-00162 Rome, Italy. E-mail: [email protected] 3Via Talamone 31/19, I-16127 Genoa. E-mail: [email protected] 4Università di Pisa, Centro interdipartimentale, Museo di Storia Naturale e del Territorio, Via Roma 79, I-56011 Calci (PI), Italy. E-mail: [email protected] 5Via Achille Zezon 10, I-20124 Milan, Italy. E-mail: [email protected] 6Dipartimento di Biologia, Università degli Studi Roma Tre, Viale Guglielmo Marconi 446, I-00146 Rome, Italy. E-mail: a_mazziotta@ hotmail.com *In: Nardi G., Whitmore D., Bardiani M., Birtele D., Mason F., Spada L. & Cerretti P. (eds), Biodiversity of Marganai and Montimannu (Sardinia). Research in the framework of the ICP Forests network. Conservazione Habitat Invertebrati, 5: 353–387. ABSTRACT A faunistic inventory of lamellicorn beetles has been conducted in southern Sardinia (former province of Cagliari) based on literature and collection records, as well as on new fi ndings from recent fi eld expeditions (2003–2008). The taxonomic analysis of the study area revealed the occurrence of 105 species (2 Lucanidae, 2 Trogidae, 7 Geotrupidae, 1 Hybosoridae and 93 Scarabaeidae). The majority of families are represented by 100% of the known Sardinian fauna; o nly the Scarabaeidae, the most diverse family, is represented by 80.8% of its Sardinian members.
  • The Phylogenetic Relationships of Tiaronthophagus N.Gen. (Coleoptera, Scarabaeidae, Onthophagini) Evaluated by Phenotypic Characters

    The Phylogenetic Relationships of Tiaronthophagus N.Gen. (Coleoptera, Scarabaeidae, Onthophagini) Evaluated by Phenotypic Characters

    insects Article The Phylogenetic Relationships of Tiaronthophagus n.gen. (Coleoptera, Scarabaeidae, Onthophagini) Evaluated by Phenotypic Characters Angela Roggero 1,* , Philippe Moretto 2, Enrico Barbero 1 and Claudia Palestrini 1 1 Department of Life Sciences and Systems Biology, University of Torino, Via Accademia Albertina 13, I-10123 Torino, Italy; [email protected] (E.B.); [email protected] (C.P.) 2 2 rue Marcel Sembat, F-83200 Toulon, France; [email protected] * Correspondence: [email protected] Received: 24 January 2019; Accepted: 20 February 2019; Published: 1 March 2019 Abstract: A necro-coprophagous new genus tha is widespread in the whole Sub-Saharan Africa was identified within the tribe Onthophagini and named Tiaronthophagus n.gen. The new genus, which is well characterized by an exclusive set of characters, comprises, at present, 26 species. Twenty species were formerly included in the genus Onthophagus and six were identified and here described as new species: Tiaronthophagus angolensis n.sp., T. jossoi n.sp., T. katanganus n.sp., T. rolandoi n.sp., T. saadaniensis n.sp., and T. zambesianus n.sp. A phylogenetic analysis that is based on a combined matrix, including discrete and landmark characters, was done. The landmark characters were tested using the geometric morphometrics techniques before their inclusion in the matrix. One single, fully resolved tree was obtained, with Tiaronthophagus constituting a distinct, monophyletic clade within Onthophagini, which was clearly separated from the other genera examined here. The biogeographical analysis identified the Central Africa as the ancestral area of the new genus and it mainly accounted for dispersal events leading to the present distribution.