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Karyology and Chromosomal Evolution of Some Small Mammals Inhabiting the Rainforest of the Rabi Oil Field, Gabon

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Karyology and Chromosomal Evolution of Some Small Mammals Inhabiting the Rainforest of the Rabi Oil Field, Gabon Karyology and Chromosomal Evolution of Some Small Mammals Inhabiting the Rainforest of the Rabi Oil Field, Gabon Ashley PRIMUS1, Jessica HARVEY1, Sylvain GUIMONDOU2, Serge MBOUMBA3, Raphaël NGANGUI4, Federico HOFFMANN5, 6, Robert BAKER5 and Calvin A. PORTER5, 1 1 Introduction no karyotypic data are available. Because some species are known to exhibit kary- The genetic information that living things inherit otypic variation within and among populations, it is from their progenitors is encoded in the molecular important to examine karyotypes from multiple indi- sequences of DNA. The DNA of animals and other viduals from different localities. Much of the kary- eukaryotes is combined with proteins and organized otypic work that has been done on small mammals in into chromosomes. Chromosomes are essential for tropical Africa has focused on Ivory Coast, the processes of cell division during mitosis and Cameroon, and Central African Republic. Some of meiosis, which distributes genetic information to Africa’s most extensive tracts of tropical rainforest daughter cells. The process of evolution often pro- are found in Gabon, and the area supports a high duces changes in the shape, number, or size of the level of biodiversity. Despite this diversity, relatively chromosomes. The karyotype of an individual is the little work has been done on the karyology of small chromosomal complement of its cells. The kary- mammals in Gabon. otype is often uniform within a species. However, in Under the auspices of the Smithsonian Institution, many cases, there is considerable karyotypic varia- and with the support of Shell Foundation and Shell tion among the populations or individuals of a Gabon, we participated in a survey of the mam- species. Comparison of karyotypes can provide valu- malian biodiversity of the Gamba Complex of able insight into relationships among species. Protected Areas. We sampled small mammals from A substantial literature has been published on the Rabi oil field, which is located in the rainforest karyotypes of small African mammals (e.g., Robbins just south of the equator in Gabon. Human activities and Baker 1978, Haiduk et al. 1980, 1981; Viegas- at Rabi are restricted to relatively small insular clear- Péquignot et al. 1983, Reumer and Meylan 1986, ings within extensive tracts of rainforest. As part of Zima et al. 1998, Schlitter et al. 1999, Biltueva et al. this survey, we prepared karyotypes from a number 2001). However, the region has a high level of bio- of species of small mammals, including shrews, logical diversity, and many species have never been rodents, and bats. We report here chromosomal data studied karyotypically. For example, the shrew genus for the shrews, rodents, and the bats of the Suborder Crocidura contains more than 150 species, of which Megachiroptera. Standard karyotypes of microchi- approximately 100 occur in Africa. Only about half of the African species have been karyotyped, and a 1 Department of Biology, Xavier University of Louisiana, considerable amount of karyotypic variation has Drexel Drive, New Orleans, LA 70125, USA. Author for been described, with diploid numbers in African correspondence, Email: [email protected] 2 Direction de la Faune et de la Chasse, BP 159, Tchibanga, Crocidura ranging from 36 to 68 (Zima et al. 1998, Gabon. Email: [email protected] Schlitter et al. 1999). Despite the lack of chromoso- 3 Gabon Biodiversity Program, Smithsonian Institution, S/C mal data for many species of Crocidura, karyotypic Shell Gabon, BP 48, Gamba, Gabon. information has been used in a number of analyses Email: [email protected] 4 IRAF - CENAREST, BP 2246, Libreville, Gabon. of evolution and systematics of this taxonomically Email: [email protected] and karyotypically diverse genus (Maddalena and 5 Department of Biological Sciences, Texas Tech University, Ruedi 1994, Zima et al. 1998, Biltueva et al. 2001). Lubbock, TX 79409, USA. Email: [email protected] 6 Although some African rodents and bats have been Present address: School of Biological Sciences, University of Nebraska, Lincoln, NE 68588. USA. well-studied, there are numerous species for which Email: [email protected] Bulletin of the Biological Society of Washington, No. 12 I 371 ropteran bats will be reported in a separate publica- Institution in Washington, or the Natural Science tion. We compare the results with published data and Research Laboratory at Texas Tech University. discuss the systematic and evolutionary implications Chromosome slides and cell suspensions are of the chromosomal variability in the taxa studied. archived with tissue samples at Texas Tech University, and are cross-referenced to the voucher 2 Materials and Methods specimens by a unique TK number. Specimens examined are listed in Table 1. We collected small mammals from the Rabi oil field of the Gamba Complex in Ogooué-Maritime 3 Results and Discussion Province, Gabon (Lee et al. this volume; see map page xxxii). Animals were sampled during February The karyotypes of 17 species (four species of insec- and March of 2002 as described by Rodriguez et al. tivores, seven species of rodents, and six of bats) (this volume) and O’Brien et al. (this volume). found in the Gamba Complex were studied, includ- Mitotic chromosome preparations were per- ing karyotypes of five species not previously docu- formed in the laboratory at Rabi from bone marrow mented: Crocidura crenata, C. goliath, C. grassei, using the methods of Baker et al. (2003), with the Heimyscus fumosus, and Scotonycteris zenkeri. For exception that ethanol was used in the fixative due to some other species, we describe karyotypes that dif- the unavailability of methanol. Meiotic preparations fer in some respects from those previously reported were made for selected males using the same proce- in the literature. Our results document that the biodi- dures with testicular tissue. Chromosomes were versity at Rabi includes chromosomal as well as tax- stained with giemsa. The Government of Gabon onomic aspects. Our field laboratory was not authorized the research and permitted retention of equipped for chromosomal banding procedures, so some specimens as vouchers. we have not yet applied chromosomal banding tech- Specimens were prepared as standard skin-and- niques to the specimens used in the study. Because skeleton or alcoholic preparations, and are deposited of this, we cannot be certain of chromosomal as vouchers in the mammal collections of the Gabon homologies across species. Standard karyotype Biodiversity Program in Gamba, the Smithsonian preparations frequently underestimate the extent of Table 1. Specimens examined with observed diploid numbers (2n). TK = unique number for cross-reference to voucher specimen. TK Species sex 2n TK Species sex 2n 110269 Crocidura crenata M 48? 110258 Hypsignathus monstrosus F36 110322 Crocidura goliath M 50 110263 Megaloglossus woermanni F34 110417 Crocidura goliath M 50 110267 Megaloglossus woermanni M34 110400 Crocidura grassei M 40 110287 Megaloglossus woermanni M34 110262 Sylvisorex ollula M 38 110290 Megaloglossus woermanni F34 110327 Sylvisorex ollula M 38 110338 Megaloglossus woermanni M34 110285 Heimyscus fumosus M 40 110264 Myonycteris torquata F36 110328 Hylomyscus parvus F 46 110265 Myonycteris torquata F36 110368 Hylomyscus stella M 46 110266 Myonycteris torquata F36 110389 Hylomyscus stella M 46 110291 Myonycteris torquata F36 110415 Lophuromys nudicaudus F 56 110292 Myonycteris torquata F36 110367 Malacomys longipes M 48 110293 Myonycteris torquata F36 110397 Mus musculoides F 34 110295 Myonycteris torquata F36 110398 Mus musculoides M 34 110299 Myonycteris torquata F36 110302 Praomys tullbergi F 34 110300 Myonycteris torquata F36 110251 Epomops franqueti F 36 110339 Myonycteris torquata F36 110253 Epomops franqueti F 36 110390 Myonycteris torquata F36 110254 Epomops franqueti F 36 110392 Myonycteris torquata M36 110463 Epomops franqueti F 36 110481 Myonycteris torquata F36 110475 Epomops franqueti M 35 110465 Scotonycteris zenkeri M32 372 I Gamba, Gabon: Biodiversity of an Equatorial African Rainforest chromosomal rearrangements (Haiduk et al. 1981, Cameroon and the Democratic Republic of Congo Baker et al. 1987). Nevertheless, our results do pro- (Hutterer and Schlitter 1996). Although the species has vide insight into karyotypic evolution in some been considered rare, Goodman and Hutterer (2004) species, and also raise some questions that can be reported a series of 13 specimens from the Monts addressed by future banding studies. Doudou region of Gabon. The karyotype of this species has not been previously reported. 3.1 Order Insectivora, Family Soricidae Chromosome preparations from a male of this shrew produced only a single spread of moderate quality (Fig. Crocidura crenata 1A). Based on this cell, the diploid number is 2n = 48. It is generally accepted that the genus Crocidura orig- The chromosomes of this cell have some banding, inated in Africa, and subsequently colonized Eurasia, although they were not treated with any banding agent. probably during the late Miocene (Maddalena and The banding was sufficient to help identify homolo- Ruedi 1994, Butler 1998). The African and Eurasian gous chromosomes, but we were unable to clearly iden- lineages both underwent extensive speciation during tify any homologies with G-banded chromosomes in the Pliocene and later. Maddalena and Ruedi (1994) other species of Crocidura (Biltueva et al. 1999, 2001). proposed a 2n = 36-40 karyotype as primitive in the We were also unable to
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