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

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