A survey of equid mitochondrial DNA: Implications for the evolution, genetic diversity and conservation of Equus

The evolution, taxonomy and conservation of the genus Equuswere investigated by examining the mitochondrial DNA sequences of thecontrol region and 12S rRNA gene. The phylogenetic analysis of thesesequences provides further evidence that the deepest node in thephylogeny of the extant species is a divergence between twolineages; one leading to the ancestor of modern horses (E.ferus, domestic and przewalskii) and the other to thezebra and ass ancestor, with the later speciation events of the zebrasand asses occurring either as one or more rapid radiations, or withextensive secondary contact after speciation. Examination of the geneticdiversity within species suggested that two of the E. hemionussubspecies (E. h. onager and E. h. kulan) onlyrecently diverged, and perhaps, are insufficiently different to beclassified as separate subspecies. The genetic divergence betweendomestic and wild forms of E. ferus (horse) and E.africanus (African ass) was no greater than expected within anequid species. In E. burchelli (plains zebra) there was anindication of mtDNA divergence between populations increasing withdistance. The implications of these results for equid conservation arediscussed and recommendations are made for conservation action.     

Homologous fission event(s) implicated for chromosomal polymorphisms among five species in the genus Equus

The genus Equus is unusual in that five of the ten extant species have documented centric fission (Robertsonian translocation) polymorphisms within their populations, namely E. hemionus onager, E. hemionus kulan, E. kiang, E. africanus somaliensis, and E. quagga burchelli. Here we report evidence that the polymorphism involves the same homologous chromosome segments in each species, and that these chromosome segments have homology to human chromosome 4 (HSA4). Bacterial artificial chromosome clones containing equine genes SMARCA5 (ECA2q21 homologue to HSA4q31. 21) and UCHL1 (ECA3q22 homologue to HSA4p13) were mapped to a single metacentric chromosome and two unpaired acrocentrics by FISH mapping for individuals possessing odd numbers of chromosomes. These data suggest that the polymorphism is either ancient and conserved within the genus or has occurred recently and independently within each species. Since these species are separated by 1-3 million years of evolution, this polymorphism is remarkable and worthy of further investigations.     

Mitochondrial genome sequence of the Tibetan wild ass (Equus kiang)

The Tibetan wild ass, or kiang (Equus kiang) is endemic to the cold and hypoxic (4000-7000?m above sea level) climates of the montane and alpine grasslands of the Tibetan Plateau. We report here the complete nucleotide sequence of the E. kiang mitochondrial genome. Our results show that E. kiang mitochondrial DNA is 16,634?bp long, and predicted to encode all the 37 genes that are typical for vertebrates.     

Polyploidy and aneuploidy inOphrys,Orchis, andAnacamptis (Orchidaceae)

Studies on chromosome numbers and karyotypes in Orchid taxa from Apulia (Italy) revealed triploid complements inOphrys tenthredinifera andOrchis italica. InO. tenthredinifera there is no significant difference between the diploid and the triploid karyotypes. The tetraploid cytotype ofAnacamptis pyramidalis forms 36 bivalents during metaphase I in embryo sac mother cells. Aneuploidy was noticed inOphrys bertolonii ×O. tarentina with chromosome numbers n = 19 and 2n = 38. There were diploid (2n = 2x = 36), tetraploid (2n = 4x = 72), hexaploid (2n = 6x = 108) and octoploid (2n = 8x = 144) cells in the ovary wall of the diploid hybridOphrys apulica ×O. bombyliflora. Evolutionary trends inOphrys andOrchis chromosomes are discussed.     

Karyotypes of three species of the family Cottidae (Scorpaeniformes)

Karyotypes and cellular DNA contents of three species of the family Cottidae viz.Icelus cataphractus, Gymnocanthus intermedius andAlcichthys alcicornis were analyzed. Structural modifications within the family were supposedly by Robertsonian translocations. The diploid chromosome numbers were determined to be 48 inAlcichthys alcicornis, 44 inGymnocanthus intermedius and 40 inIcelus cataphractus. The DNA contents ranged from 1.46 to 1.50pg/cell in the three species. The karyotype ofIcelus cataphractus is unique in having the smallest chromosome number (2n = 40) and 14 large-sized chromosomes. From the chromosome number and the existence of some large chromosome pairs, Robertsonian translocations seem to have occurred frequently inIcelus cataphractus andGymnocanthus intermedius.     

Chromosomal polymorphism in Equus hemionus.

The first cytogenetic studies of the Turkmenian kulan, Equus hemionus kulan, are reported, and a polymorphism in diploid chromosome number is described. Chromosome fusion is apparently involved in the alterations of the karyotype of E. hemionus kulan (2n = 55, 54) when compared to the karyotype of the onager, E. hemionus onager (2n = 56). Additionally, the rearrangement involved has been identified in animals unrelated through captive breeding; inheritance of the fusion chromosomes has also been observed.     

Distinction in morphology and esterase isozyme betweenEupatorium glehni (=E. chinense subsp.sachalinense) andE. chinense var.oppositifolium (compositae)

The taxonomic status ofEupatorium chinese var.oppositofolium andE. glehni (=E. chinense subsp.sachalinense) inE. chinense complex semsu Kitamura has long been controversial. In this paper, the degree of divergence between diploids of these two taxa was examined by means of morphological studies including principal component analysis, the electrophoretic analysis of esterase isozyme variation and observations on habitats. The data obtained through the examinations indicate these two taxa are diverged enough to be recognized as distinct biological species. Since the polyploidE. chinense var.oppositifolium is more or less intermediate in morphology between the two diploid taxa, it is considered to have masked the distinction between the two diploid taxa. Also, electrophoretic evidence suggests that polyploidE. chinense var.oppositifolium is not a hybrid or hybrid derivative withE. glehni as a parental species. Possible origin of polyploidE. chinense varoppositifolium is also discussed.     

Chromosome numbers, karyotypes and nuclear DNA contents from perennial polyploid groups ofCerastium (Caryophyllaceae)

Somatic chromosome numbers have been determined for the followingCerastium taxa:C. eriophorum (2n = 36),C. alpinum (2n = 72),C. transsylvanicum (2n = 108),C. arcticum (2n = 108),C. latifolium (2n = 36),C. carinthiacum (2n = 36),C. banaticum (2n = 36),C. arvense subsp.glandulosum (2n = 36),C. arvense subsp.arvense (2n = 72) andC. fontanum (2n = 144). Karyotypes of three diploid species (C. eriophorum, C. banaticum andC. latifolium), belonging to three different taxonomic groups, were analysed and found to be similar. The relative nuclear DNA contents of all taxa were determined by flow cytometry and, for five species, also by Feulgen cytophotometry. The values obtained by the two methods are similar. A comparison of nuclear DNA contents among diploids shows that values differ significantly between different taxonomic groups, and are correlated with average chromosome size. Within closely related polyploid groups nuclear DNA amounts increase from 2x- to 4x- and 6x taxa as 1 : 1.4 : 2.4 in theC. alpinum complex, whereas DNA amounts are doubled comparing 2x- and 4x-subspecies in theC. arvense complex.     

马科在中国的分布与现状

蒙古野驴(Asinus h．hemionus)分布于内蒙和甘肃西北角、新疆东北部至准噶尔盆地西部。1982年航调,卡拉麦里山保护区内有356头,O.02头／平方公里;1986年调查为O.03头／平方公里。估计目前我国境内不超过2000头。西藏野驴(Asinus k.kiang)分布于西藏西部、新疆南缘山地;A.k.holdereri分布于除湟水河谷外青海全境、四川西北角、甘肃祁连山。1984年调查,阿尔金山保护区内有41,262头,0.917头／平方公里。估计我国境内种群约在20万头上。多年考察未发现野马(Equus przewalskii),目前可能在北塔山以东尚有少数残存。     

Variation in wild populations of annual beet (Beta,Chenopodiaceae)

Among the morphological variation in wild annual populations of sect.Beta only tepal characters show a geographic pattern, and hence can be used to distinguish different taxa. Two morphological types correspond with taxa already described:B. macrocarpa (incl.B. bourgaei) has a Macaronesian and W. to E. Mediterranean andB. adanensis an E. Mediterranean distribution area. A third type in the Aegean region is not well known yet and possibly has to be included inB. macrocarpa. Both diploid and tetraploid (x = 9) cytotypes are found withinB. macrocarpa, the latter exclusively on the Canary Islands.     

Spontaneous triploidy and NOR activity in Eigenmannia sp. (Pisces,Sternopygidae) from the Amazon basin

The occurrence of a natural triploid among diploid individuals of Eigenmannia sp from the Amazon basin is described. The diploid individuals had 2n=31 and 32 chromosomes, the difference in chromosome number being due to a Robertsonian rearrangement. The triploid individual presented 2n=46 and had two fused acrocentrics. NOR activity was present in the three homologues of triploid and a NOR polymorphism was detected in the triploid and diploids.     

Mitochondrial DNA evolution in the genus Equus

Employing mitochondrial DNA (mtDNA) restriction-endonuclease maps as the basis of comparison, we have investigated the evolutionary affinities of the seven species generally recognized as the genus Equus. Individual species' cleavage maps contained an average of 60 cleavage sites for 16 enzymes, of which 29 were invariant for all species. Based on an average divergence rate of 2%/Myr, the variation between species supports a divergence of extant lineages from a common ancestor approximately 3.9 Myr before the present. Comparisons of cleavage maps between Equus przewalskii (Mongolian wild horse) and E. caballus (domestic horse) yielded estimates of nucleotide sequence divergence ranging from 0.27% to 0.41%. This range was due to intraspecific variation, which was noted only for E. caballus. For pairwise comparisons within this family, estimates of sequence divergence ranged from 0% (E. hemionus onager vs. E. h. kulan) to 7.8% (E. przewalskii vs. E. h. onager). Trees constructed according to the parsimony principle, on the basis of 31 phylogenetically informative restriction sites, indicate that the three extant zebra species represent a monophyletic group with E. grevyi and E. burchelli antiquorum diverging most recently. The phylogenetic relationships of E. africanus and E. hemionus remain enigmatic on the basis of the mtDNA analysis, although a recent divergence is unsupported.      

Chromosomal polymorphism associated with Robertsonian fusion in Seriola dumerili (Risso, 1810) (Pisces: Carangidae)

The diploid numbers 2 n = 48, and 2 n = 47 have been determined for the greater amberjack, Seriola dumerili. A chromosome polymorphism due to Robertsonian fusion is present in this species. A simple sex-determining mechanism has not been observed.     

Chromosomal variation in the southern short-tailed shrew (Blarina carolinensis)

Chromosomal polymorphism was assessed in the southern short-tailed shrew (Blarina carolinensis) using standard metaphase chromosome and G-banding techniques. Twenty-one animals (11 males, 10 females) from the Meeman Biological Station in Shelby Co., Tennessee, were examined for diploid number. Results showed diploid numbers of 35, 36, 37, 38, 39, 40 and 41 and fundamental numbers of 41, 42, 43, 44 and 45. No diploid numbers or fundamental numbers were unique to a specific collecting locality. The first G-banded karyotypes are reported for the species. These results indicate that Robertsonian polymorphisms, inversions, and possibly other events are responsible for chromosomal variation in B. carolinensis.     

Ancient DNA from Nubian and Somali wild ass provides insights into donkey ancestry and domestication

Genetic data from extant donkeys (Equus asinus) have revealed two distinct mitochondrial DNA haplogroups, suggestive of two separate domestication events in northeast Africa about 5000 years ago. Without distinct phylogeographic structure in domestic donkey haplogroups and with little information on the genetic makeup of the ancestral African wild ass, however, it has been difficult to identify wild ancestors and geographical origins for the domestic mitochondrial clades. Our analysis of ancient archaeological and historic museum samples provides the first genetic information on the historic Nubian wild ass (Equus africanus africanus), Somali wild ass (Equus africanus somaliensis) and ancient donkey. The results demonstrate that the Nubian wild ass was an ancestor of the first donkey haplogroup. In contrast, the Somali wild ass has considerable mitochondrial divergence from the Nubian wild ass and domestic donkeys. These findings resolve the long-standing issue of the role of the Nubian wild ass in the domestication of the donkey, but raise new questions regarding the second ancestor for the donkey. Our results illustrate the complexity of animal domestication, and have conservation implications for critically endangered Nubian and Somali wild ass.     

Cytotaxonomy and breeding system of the genusBiscutella (Cruciferae)

Chromosome counts were determined for 46 populations ofBiscutella representing 28 taxa. The genus was found to contain diploid taxa with 2n = 12, 16 and 18, tetraploid taxa with 2n = 36 and hexaploid taxa having 2n = 54.B. laevigata L. s. l. consists of diploid and tetraploid populations which are poorly differentiated morphologically. TetraploidB. laevigata s. l. and hexaploidB. variegata Boiss. & Reuter (s. l.) are characterized by chromosomal instability. The variation in chromosome numbers and the occurrence of polyploidy is discussed in relation to the taxonomy of the genus. An investigation of the breeding system showed that most of the annual species were self-compatible and partly inbreeding and most of the perennial species self-incompatible and, therefore, outbreeding, while one annual species,B. cichoriifolia Loisel., showed both systems.     

月季种质资源的倍性变异及核型多样性研究(英文)

采用常规压片法对包括5个蔷薇野生种、8个古老月季及9个现代月季品种共22份月季种质资源进行核型分析。供试材料共有3种不同倍性:5个野生种中粉团蔷薇为三倍体(2n=3x=21),其他4个材料为二倍体(2n=2x=14);9个现代月季品种均为四倍体(2n=4x=28);而古老月季品种中包含二、三、四倍体3种倍性。22份材料中共发现4种不同类型的核型,在野生种中为1A、2A;古老月季中为1A、2A、1B;现代月季中为1A、1B、2B。另外,供试材料在核不对称系数、着丝粒指数、随体数目及位置、染色体相对长度组成等方面差异较大,说明在月季种质资源中存在着丰富的核型多样性。     

New Ctenomys karyotypes (Rodentia,Octodontidae) from north-eastern Argentina and from Paraguay confirm the extreme chromosomal multiformity of the genus

Bone-marrow karyotypes of 68 specimens of the subterranean octodontid rodent genus Ctenomys from 16 different populations of north east Argentina and one from Paraguay have been studied. A surprising variety of chromosome numbers was found, ranging from 2n=42 to 2n=70. Some of the karyomorphs are clearly assigned to named species by topotypy: C. conoveris 2n=50, FN=56; C. argentinus, 2n=44, FN=54; C. perrensi, 2n=50, FN=84; C. dorbignyi, 2n=70, FN=84; C. roigi, 2n=48, FN=80; C. yolandae, 2n=50, FN=78. Four populations of Corrientes Province similar in morphology to C. perrensi were found to be polymorphic and polytypic; they maintain the same FN=84, but diploid numbers increase from 2n=54 to 2n=58 from SW to the NE, thus suggesting Robertsonian rearrangements. In the middle of this cline, a stable karyomorph of 2n=62, FN=84 was found in two different populations, suggesting to belong to an undescribed species. Another karyomorph of 2n=42, FN=76 found in Curuzú Laurel, Corrientes, may also prove to represent another undescribed species. One karyomorph of 2n=52, FN=74, and another of 2n=56, FN=78 from Paraná and Ubajaý (Entre Ríos Province, Argentina) respectively are close to C. rionegrensis. The relationships among these karyomorphs is considered in light of data on sperm morphology. The hypothesis is advanced that karyotypic rearrangements among the FN=84 group may be the result of Robertsonian repatterning from a 2n=70 original widespread form. Fixation of chromosomal variants is correlated with patchy distribution and small size of unstable demes, and may or may not have resulted in reproductive isolation.     

A complex chromosomal polymorphism in Gobius fallax (Gobiidae,Perciformes)

Chromosome analysis of 53 specimens from a population of Gobius fallax has revealed inter- and intra-individual variation in the diploid number (2n=38–43) arising mainly through Robertsonian translocations. The presence of a small biarmed chromosome in a few cells from 3 males as well as the numerous multivalent configurations in meiotic plates and the apparent association between the NOR-chromosomes and other acrocentrics suggest that this polymorphism has an inherited as well as a somatic origin and that some translocations may involve more than two acrocentric pairs simultaneously.     

Karyosystematic studies on threeCrepis species (Asteraceae) endemic to Greece

This work examines the cytogeographical distribution, the morphological characters, and the karyotypes of threeCrepis species endemic to Greece (C. sibthorpiana, C. incana, andC. heldreichiana). C. sibthorpiana is diploid (2n = 2x = 8),C. incana is diploid (2n = 2x = 8) and tetraploid (2n = 4x = 16, 17), andC. heldreichiana is always dekaploid (2n = 10x = 40). The Giemsa positive bands, usually pairs of dots, are mainly centromeric inC. incana, while they are terminal inC. sibthorpiana (on the short arm of all chromosomes) and inC. heldreichiana (on both arms of all chromosomes). Intercalary C-bands are scarce and usually variable within karyotypes, individuals, and species. The most variable karyotype both in Feulgen and Giemsa preparations is that ofC. incana, in which also supernumerary chromosomes were observed, which are polysomic to standard set members. On the basis of morphological and karyological data the evolutionary relationships among the threeCrepis taxa are discussed.