Genetic diversity of mitochondrial cytochrome b gene in swamp buffalo

The swamp buffalo (Bubalus carabanensis) is mainly bred for meat, transport and rice cultivation in China and Southeast Asian countries. In the current study, we investigated the genetic diversity, maternal origin and phylogenetic relationship of swamp buffalo by analyzing 1,786 mitochondrial cytochrome b (cytb) sequences from China, Vietnam, Laos, Thailand, India and Bangladesh. Our results indicated that the swamp buffalo can be divided into two major lineages (SA and SB with the sublineages) and three rare lineages (SC, SD and SE), which showed strong geographic differentiation. The SA1 lineage represented a major domestication event, which involved population expansion. Regions III and V showed higher haplotype diversity than the other regions, indicating that the regions of Southwest China and IndoChina are potential domestication centers for the swamp buffalo. In addition, the swamp buffalo showed a closer phylogenetic relationship to tamaraw. In conclusion, our findings revealed a high level of genetic diversity and the phylogenetic pattern of the swamp buffalo.     

Genetic diversity of Asian water buffalo (Bubalus bubalis): mitochondrial DNA D-loop and cytochrome b sequence variation

Swamp and river buffalo mitochondrial DNA (mtDNA) was sequenced for 303 bp of the cytochrome b gene for 54 animals from 14 populations, and for 158 bp of the D-loop region for 80 animals from 11 populations. Only one cytochrome b haplotype was found in river buffalo. Of the four haplotypes identified in swamp buffalo, one found in all populations is apparently ancestral both to the other swamp haplotypes and to the river haplotype. The phylogenetic relationships among the 33 D-loop haplotypes, with a cluster of 11 found in swamp buffalo only, also support the evolution of domesticated swamp and river buffalo from an ancestral swamp-like animal, most likely represented today by the wild Asian buffalo ( Bubalus arnee ). The time of divergence of the swamp and river types, estimated from the D-loop data, is 28 000 to 87 000 years ago. We hypothesise that the species originated in mainland south-east Asia, and that it spread north to China and west to the Indian subcontinent, where the rive type evolved and was domesticated. Following domestication in China, the domesticated swamp buffalo spread through two separate routes, through Taiwan and the Philippines to the eastern islands of Borneo and Sulawesi, and south through mainland southeast Asia and then to the western islands of Indonesia.     

Analysis of genetic diversity of domestic water buffaloes and anoas based on variations in the mitochondrial gene for cytochrome b

There are two major groups of domestic water buffaloes in East and Southeast Asia: swamp buffaloes and river buffaloes. Genetic diversity among swamp and river buffaloes was studied by DNA sequence analysis of the mitochondrial gene for cytochrome b. The results showed that each of the two groups has mitochondrial DNA (mtDNA) with a specific cytochrome b haplotype. The pairwise nucleotide sequence divergence was calculated to be 2·67% between swamp and river buffaloes, suggesting that they might have diverged from the ancestral populations of Asiatic domestic water buffaloes, approximately 1 million years ago. In addition, the sequences of the same gene from three subspecies of anoa (lowland, mountain and quarles anoa) were determined and compared with that of a domestic water buffalo. The sequence divergence was 1·2% for mountain anoa vs quarles anoa, 3·6% for mountain anoa vs lowland anoa and 3·3% for quarles anoa vs lowland anoa. Moreover, the sequence divergence between water buffaloes and anoas was found to be approximately 3·33%. Our results provide molecular evidence to support the taxonomic classification, namely, that Asiatic buffaloes may be classified into four lineages, swamp buffalo, river buffalo, lowland anoa and mountain plus quarles anoa. However, the sequence divergence values among these four groups were lower than the sequence divergence values found in the genus and subgenus levels within the subfamily Bovinae. In particular, in contrast to some proposed taxonomic classifications, our results indicated that mtDNA in the water buffaloes and anoas did not diverge at the genus level.     

Characterization of cytochrome b diversity in Chinese domestic horses

Previous mitochondrial DNA (mtDNA) D‐loop and microsatellite studies have shown that Chinese horses have multiple maternal origins and high genetic diversity. To better characterize maternal genetic origins and diversity of Chinese domestic horses, we conducted a comprehensive analysis of 407 complete 1140 bp sequences of the horse mitochondrially encoded cytochrome b (CYTB) gene, including 323 horses from 13 Chinese indigenous breeds and 84 reference sequences from GenBank. A total of 114 haplotypes were identified, of which 73 appeared among the 13 Chinese horse breeds. The high mitochondrially encoded cytochrome b haplotypic diversity suggests multiple maternal origins in Chinese horses.     

大鳍鱊基于细胞色素b基因序列的遗传变异及生物地理过程

对采集于长江水系、黑龙江水系及海南岛的36尾大鳍(Acheilognathus macropterus)的细胞色素b基因序列进行了研究分析。发现有须个体和无须个体在分子系统树中处于混合分布状态,各个体相互间遗传变异远小于与外类群(A.chankaensis、A.tokinensis)间差异,统计了其主要形态度量数据,也显示各主要性状数据无显著差异,结果均支持有须个体和无须个体(曾经被鉴定为A.taenianalis)为同一物种,A.taenianalis为A.macropterus的同物异名。序列分析显示,36个个体明显聚合成5个大的分支,其中clade1分支全部由黑龙江水系类群组成,clade5分支由海南岛的1个个体组成,长江水系的所有个体分散在clade2、clade3、clade4这三个分支中,显示出一定的地理相关性。在clade2、3、4中,clade2包括了长江上游的5个个体和长江中游的2个个体,同时还包括了黑龙江的2个个体;clade3包括了中游的2个个体;clade4包括了中游的12个个体,三个分支间又存在明显的遗传变异率(均>5%),从21个个体在三个分支中的分布情况来看,三者间这种较大的遗传变异率不能由地理隔离解释,故推测,造成同一水系的A.macropterus间出现较明显遗传分化的原因可能与其特殊的性选择生殖方式有关,该生殖方式限制了不同生殖集群间同种鱼类的基因交流。基于系统分支图,计算了各分支间的遗传距离,由结果推测大鳍的生物地理过程为:海南岛水系类群同大陆类群较早产生隔离,黑龙江水系类群由长江水系类群在晚近时期向北扩散形成。       

Genetic diversity and the biogeographical process of Acheilognathus macropterus revealed by sequence variations of mitochondrial cytochrome b gene

In this study, thirty-six individuals of Acheilognathus macropterus were collected from the Heilongjiang River,the Yangtze River,and the Nandujiang River.Partial mitochondrial cytochrome b gene region (636 base pair) was sequenced to these samples and 22 haplotypes were found.With A.chankaensis and A.tokinensis as outgroups,their relationships were analyzed.The p-distances were calculated with Mega software and a molecular phyiogenetic tree was constructed using the neighbor-joining (NJ) method.The proportions of main morphological characters were compared as well.P-distances showed that the genetic differences in A.macropterus samples were far smaller than those between these samples and the outgroups.The molecular phylogenetic tree shows that samples with barbels and those without barbels were intermingled.There was no distinctive difference in proportions of morphological characteristics among them.These results suggested that samples with barbels and those without barbels (formally identified as A.taenianalis) are the same species;A.taenianalis is synonymous with A.macropterus.The thirtysix individuals were grouped into five clades and the positions of the samples in the clades were correspondingly grouped within their geographical distributions.Among the five clades,clades 1 and 5 included samples from the Heilongjiang River and Nandujiang River respectively.The samples from the Yangtze River scattered into clades 2,3,and 4.There were distinctive genetic differences (> 5%)among them.Interestingly,the distributions of the 21 samples in these three clades were not correlated to their geographical distributions.It is postulated that these genetic differences were due to the bitterlings' mating choice mechanism,the prozygotic isolation.The genetic differences between the fish from Nandujiang River and those from the mainland indicated that they were separated early.However,the small genetic differences among the samples and the positions of the fish from the Heilonjiang River in the molecular phylogenetic tree indicate that fish in Heilongjiang River might have dispersed from the Yangtze River to that area much later.     

Genetic diversity and differentiation of Chinese domestic buffalo based on 30 microsatellite markers

To determine genetic diversity and evolutionary relationships among Chinese buffalo populations, 18 indigenous swamp buffalo populations and two introduced river buffalo breeds were genotyped for 30 microsatellite loci. The mean number of alleles across the 30 loci was 8.13, and the expected heterozygosity ranged from 0.517 (Yanjin) to 0.609 (Dehong). Although there was abundant genetic variation, genetic differentiation between Chinese buffalo populations was low, with only 2.8% of the total genetic variance among populations. The genetic differentiation pattern and genetic relationships among Chinese buffalo populations were consistent with their geographic distribution. The Dehong population was discerned as a distinct indigenous population, but suffered genetic admixture with river buffalo. The remaining populations were divided into four major clusters, i.e. the Upper and Middle Reaches of Yangtze Valley cluster (Guizhou, Guizhoubai, Yanjin, Fuling, Enshi and Jianghan), the Lower Reaches of Yangtze Valley cluster (Haizi, Shanqu and Dongliu), the South China cluster (Fuan and Xinfeng) and the Southwest China cluster (Xinglong, Xilin, Diandongnan and Dechang).     

斑鳠细胞色素b基因的初步研究

斑鳠是珠江四大名鱼之一,目前资源量急剧下降,种质资源的保护迫在眉睫。测定了斑鳠线粒体细胞色素b基因的全序列,该序列全长1137bp,其中T、C、A、G四种碱基含量分别为31.6%、25.7%、29.6%、13.1%,A+T的含量(61.2%)显著高于G+C的含量(38.8%)。将测定序列(序列自编号:MGGD)与GenBank中的2个斑鳠序列(序列自编号:MGZJ、MGFJ)进行了比较,3个序列之间的序列差异为0.005-0.017,转换后的氨基酸序列间差异位点都仅为2个,反应出细胞色素b基因作为功能蛋白质编码基因序列相对比较保守。鳠属鱼类线粒体Cytb基因的转换大于颠换。研究为斑鳠渔业种质资源保护和品种选育提供了遗传方面的资料,斑鳠的群体结构有待进一步深入研究。     

中国地方黄牛的遗传多样性*

中国黄牛起源复杂,我国地方黄牛群体不同品种在毛色、形态外貌、细胞遗传学、血液蛋白座位分析均表现出多样性。计算我国黄牛群体6个毛色座位平均杂合度和6个血液蛋白座位平均杂合度分别为0.3144和0.4873,表明我国地方黄牛群体的遗传多样性非常丰富。计算我国黄牛群体的6个毛色座位和6个血液蛋白座位的基因分化系数分别为0.3404和0.095,表明我国黄牛群体毛色差异中有34.04％是由品种间的差异造成的。血液蛋白的多态性有9.5％是由品种间的差异造成的。我国黄牛群体的遗传多样性主要来自品种内的遗传多样性。保存我国黄牛品种资源多样性不仅要从整个中国黄牛群体上考虑,而且要针对每个品种（或类群）进行保种。     

基于线粒体细胞色素b基因的中国大沙鼠系统地理格局

通过内蒙、新疆、甘肃的41个大沙鼠样品和1个伊朗撒拉克大沙鼠的mtDNA Cytb基因全序列的遗传分析,对我国大沙鼠(Rhombomys opimus)的分子系统地理学进行了初步探讨。结果表明,我国41个大沙鼠样品的Cytb基因包含了50个核苷酸变异位点(占全序列的4.39%),其中转换48个,颠换2个,共定义23个单倍型。在四个地理种群中,内蒙古中部半荒漠区和阿拉善荒漠区的单倍型多样性最高,甘新荒漠区的单倍型多样性最低;北疆荒漠区的核苷酸多样性最高,内蒙古中部半荒漠区的核苷酸多样性最低。分子变异分析(AMOVA)表明,种群间的遗传变异占51.68%,种群内的遗传变异占48.32%。FST统计结果表明,除内蒙古中部半荒漠区与阿拉善荒漠区地理种群之间差异显著外(P<0.05),其它地理种群间的差异均极显著(P<0.01)。基于单倍型的系统树显示,42只大沙鼠形成三支。其中,伊朗撒拉克地区大沙鼠和中国地区大沙鼠之间的亲缘关系比中国两支大沙鼠之间的亲缘关系远;分析表明,中国分布的大沙鼠两支之间分歧时间估计在0.093Ma前。嵌套支分析表明,大沙鼠历史种群曾发生过异域片段化、受阻碍基因流和持续种群扩张事件。种群扩张分析提示大沙鼠在0.0119Ma前曾经历过一次种群扩张事件,种群可能受到末次冰期波动的影响。     

翘嘴鳜线粒体DNA细胞色素b基因序列分析

测定了翘嘴鳜（Siniperca chuatsi）的线粒体细胞色素b基因编码区全序列1140个核苷酸序列,并且由此推导出对应的氨基酸序列,A/T所占比例为53.4％,G C约占46.6％。该基因中密码子第1位核苷酸中4种碱基组成较为均衡,第2位核苷酸中T的使用率较高,G的使用率较低;而密码子第3位C/A的使用率高,而G的使用率仅为3.7％;在氨基酸序列中Leu所占比例16.36％,远高于其它氨基酸;而Cys使用比例仅为0.79％。本研究为鳜类的系统发育,资源保护和利用提供了核苷酸序列方面的资料,至于翘嘴鳜线粒体细胞色素b基本序列中密码子不同位置碱基使用比率以及氨基酸组成是翘嘴鳜特有还是鳜类共有特征,尚有待进一步研究。     

昆明城郊中国树鼩群体线粒体DNA遗传多样性

由于树鼩是灵长类动物的近亲,且具有体型小、繁殖周期短、饲养管理成本低等优点,长期以来被认为有望替代灵长类动物用于人类疾病的动物模型研究.然而,目前对树鼩的群体遗传结构还知之甚少,这极大地限制了其在疾病动物模型研究的应用,也是其品系资源创制的瓶颈.本研究通过分析80只采自于云南省昆明周边地区的野生树鼩(Tupaia belangeri chinensis)线粒体DNA(mtDNA)多态性,结合国外报道的2个树鼩(Tupaia belangeri)序列比较后发现,在604 bp的mtDNA控制区片段中兵检测到29个核苷酸替代变异,这些变异共界定了13种单倍型,表现较高的群体遗传多样度.另外,昆明地区的树鼩与国外报道的2个树鼩间存在较大的遗传分化,mtDNA控制区单倍型之间的核苷酸替换数大于18个,远高于昆明地区树鼢群体内部不同单倍型之间的差异.选择含有代表性的mtDNA控制区单倍型的17个昆明地区树鼩个体进一步测定了细胞色素b基因片段(1134 bp),结合前人报道的数据分析,结果进一步支持mtDNA控制区数据反映的遗传格局及揭示的昆明地区树询与国外报道树鼩之间的明显差异.本研究结果提示,昆明地区树鼩与国外树鼩之间存在较大遗传差异,在将树鼩用于人类疾病动物模型研究中要注意这些遗传差别.昆明城郊的树鼩群体具有较高的遗传多样度,在开展近交系建立等工作时须考虑选取群体内部具有代表性的mtDNA世系.

Abstract：

Due to their special phylogenetic position in the Euarchontoglires and close affinity to primates, tree shrews have been proposed as an alternative experimental animal to primates in biomedical research. However, the population genetic structure of tree shrews has largely remained unknown and this has hindered the development of tree shrew breeding and selection. Here we sampled 80 Chinese tree shrews (Tupaia belangeri chinensis) in Kunming, China, and analyzed partial mtDNA control region sequence variation. Based on our samples and two published sequences from northern tree shrews (T. belangeri), we identified 29 substitutions in the mtDNA control region fragment (～ 604 bp)across 82 individuals and defined 13 hapiotypes. Seventeen samples were selected for sequencing of the cytochrome b (Cyt b; 1134 bp) gene based on control region sequence variation and were analyzed in combination with 34 published sequences to solidify the phylogenetic pattern obtained from control region data. Overall, tree shrews from Kunming have high genetic diversity and present a remarkable long genetic distance to the two reported northern tree shrews outside China. Our results provide some caution when using tree shrews to establish animal models because of this apparent genetic difference. In addition, the high genetic diversity of Chinese tree shrews inhabiting Kunming suggests that systematic genetic investigations should be conducted before establishing an inbred strain for medical and biological research.     

基于细胞色素b序列的鲹科分子系统发育

测定了中国鲹科8属9种鱼的细胞色素b基因的全序列(1141bp),结合来自GenBank中分布于美国、安哥拉、希腊以及巴拿马的鲹科4属14种鱼的相应同源序列生成供系统发育分析的序列矩阵,用最大简约法和邻接法构建分子系统树。结果显示:(1)支持科下设四个亚科(鲹亚科,亚科,鲳鲹亚科,鰆鲹科)阶元的分类系统;(2)亚科属下不宜设亚属分类阶元;(3)及达副叶鲹与丽叶鲹亲缘关系较近,应同属于副叶鲹属;(4)我国传统的鱼类检索系统将狮鼻鲳鲹误鉴定为卵形鲳鲹,建议予以修正。     

鳜类系统发育的线粒体Cytb基因全序列分析

测定了鳜、大眼鳜、斑鳜、暗鳜、波纹鳜、长体鳜、中国少鳞鳜等7种鳜类12个个体的线粒体细胞色素b基因全序列。结合GenBank中的同源序列,共分析了9种鳜类的系统发育关系。序列分析表明,鳜属鱼类属内种间的遗传距离(0.015～0.093)明显小于少鳞鳜属鱼类属内种间的遗传距离(0.152～0.178)。在分子系统发育树上,长体鳜与鳜属的鳜、大眼鳜、斑鳜、波纹鳜、暗鳜聚合成一分支,少鳞鳜属的种类聚成另一分支;支持将长体鳜归入鳜属,鳜类分为鳜属和少鳞鳜属等二个属的分类处理。在鳜属鱼类中,鳜和大眼鳜亲缘关系十分密切;斑鳜与波纹鳜亲缘较近;长体鳜与鳜属其它5个种的亲缘关系较远。在少鳞鳜属鱼类中,中国少鳞鳜和日本少鳞鳜的亲缘关系较远,韩国少鳞鳜的系统位置较不明确。鳜类的单系性及其鳜类的系统位置仍有待进一步研究。     

Site heteroplasmy in the mitochondrial cytochrome b gene of the sterlet sturgeon Acipenser ruthenus

Sturgeons are fish species with a complex biology. They are also characterized by complex aspects including polyploidization and easiness of hybridization. As with most of the Ponto-Caspian sturgeons, the populations of Acipenser ruthenus from the Danube have declined drastically during the last decades. This is the first report on mitochondrial point heteroplasmy in the cytochrome b gene of this species. The 1141 bp sequence of the cytb gene in wild sterlet sturgeon individuals from the Lower Danube was determined, and site heteroplasmy evidenced in three of the 30 specimens collected. Two nucleotide sequences were identified in these heteroplasmic individuals. The majority of the heteroplasmic sites are synonymous and do not modify the sequence of amino acids in cytochrome B protein. To date, several cases of point heteroplasmy have been reported in animals, mostly due to paternal leakage of mtDNA. The presence of specific point heteroplasmic sites might be interesting for a possible correlation with genetically distinct groups in the Danube River.     

A salmon id phylogeny inferred from mitochondrial cytochrome b gene sequences

Partial mitochondrial cytochrome b gene sequences of eight salmonid species were used in a PAUP analysis to generate a phylogeny of the group. The four genera represented are Salmo, Salvelinus, Oncorhynchus and Thymallus . The inferred phylogenetic tree coincides well with the classically derived one for these genera. The recent reclassification of the rainbow trout as a member of the genus Oncorhynchus is supported. The assignment of grayling as the outgroup is vindicated. The utility of gene sequence data to infer the phylogenetic relationships of the Salmonidae is discussed.     

大林姬鼠两亚种线粒体DNA 细胞色素b 基因和控制区的序列多样性

为了研究大林姬鼠两亚种(韩国的指名亚种及中国东北和内蒙古地区的东北亚种)线粒体DNA 的变异程度并确定朝鲜亚种的分类地位,我们分别将来自韩国和中国东北长白山地区的两亚种的线粒体DNA 的细胞色素b 基因和控制区进行了测序分析。我们将测序所得到细胞色素b 基因序列与来自基因库的大林姬鼠5 个亚种的相应的单倍型进行了分析,结果显示大林姬鼠可分为4 个类群[类群1:韩国大林姬鼠指名亚种;类群2:中国长白山和内蒙古地区的东北亚种、俄罗斯外加贝尔的majuculus 亚种;类群3:中国长春的东北亚种、俄罗斯Primorye(俄罗斯远东地区) rufulus 亚种、俄罗斯库页岛(俄罗斯远东地区)和日本北海道地区的giliacus 亚种;类群4:中国黑龙江海林地区的东北亚种]。线粒体的控制区序列分析显示韩国指名亚种也不同于中国东北地区的东北亚种。本研究的类群1,2 和3 与Serizawa et al. (2002)的研究的K、S 和R 的分支相对应。这表明韩国指名亚种(类群1 和分支K)的线粒体DNA 与其他类群不同。另外,我们还发现在细胞色素b 基因构建的系统树中,东北亚种可以与类群2 (分支S)及类群3 (分支R 的不同亚种聚合在一起。我们认为线粒体DNA 的母性遗传与两个相邻亚种的个体之间的种内杂交造成了基于细胞色素b 序列对东北亚种的聚类分析结果与基于形态学特征的分类结果的不一致。因此,我们提出对这些显示出核苷酸序列多样性的东北亚种不能只用细胞色素b的数据进行亚种分类,还应该结合形态学和核DNA 特征进行进一步分析。最后,我们还发现韩国的指名亚种的细胞色素b 序列在平均距离16. 93% 的基础上不同于来自基因库的A. speciosus。Jones and Johnson (1965)指出了韩国的大林姬鼠在形态上的区别,所以我们认为韩国的大林姬鼠指名亚种A. p. peninsulae 是一种具有形态和遗传特异性的地方亚种。     

Genetic structure of Mediterranean populations of the sandfly Phlebotomus papatasi by mitochondrial cytochrome b haplotype analysis

Phlebotomus papatasi (Scopoli) (Diptera: Psychodidae) is the main vector of Leishmania major Yakimoff & Schokhor; which is the cause of self-limiting cutaneous leishmaniasis in the Old World. This sandfly is found in houses, animal shelters, caves and rodent burrows. It has a large geographical range, which includes the Middle East and the Mediterranean regions. A population analysis of colony and field specimens of P. papatasi was conducted on 25 populations originating from 10 countries. The distribution of haplotypes of the maternally inherited mitochondrial cytochrome b gene were analysed to assess the population differentiation of P. papatasi. Alignment of a 442-basepair region at the 3' end of the gene identified 21 haplotypes and 33 segregating sites from 131 sandflies. The pattern of sequence variations did not support the existence of a species complex. The median-joining network method was used to describe both the origin of the haplotypes and the population structure; haplotypes tended to cluster by geographical location, suggesting some level of genetic differentiation between populations. Our findings indicate the presence of significant population differentiation for populations derived from Syria, Turkey, Palestine, Israel, Jordan and Egypt. Knowledge of population differentiation among P. papatasi populations is important for understanding patterns of dispersal in this species and for planning appropriate control measures.     

PCR-RFLP analysis of the cytochrome b gene in horse mitochondrial DNA

The mitochondrial DNA sequence of cytochrome b gene in a Thoroughbred horse was determined. By comparing DNA sequences between the Thoroughbred and published sequence data (two horses and one Grevyi zebra), polymerase chain reaction (PCR) primers were designed for amplification of a 590 bp DNA fragment in the cytochrome b gene, and PCR-restriction fragment length polymorphism (RFLP) analysis was studied in 140 horses of six breeds using three restriction enzymes ( AciI, BamHI, RsaI ). Two morphs were found using each of the three enzymes. By combining three enzymes morphs, the 140 horses examined were classified into four types. Type 2 was most frequent in all breeds.     

Genetic diversity and historical population structure in the New Zealand mayfly Acanthophlebia cruentata

1. Nucleotide sequences of a 280 base pair region of the cytochrome b gene were used to assess genetic diversity and to infer population histories in the New Zealand mayfly Acanthophlebia cruentata. 2. A hierarchial examination of populations from 19 streams at different spatial scales in the central and northern North Island of New Zealand found 34 haplotypes. A common haplotype was found in all central region streams and unique haplotypes in northern streams. Several central streams had region specific haplotypes with genetically differentiated populations at the 70–100 km scale. 3. Haplotype diversity was high (0.53–0.8) at most sites, but low (0–0.22) in some central sites. amova analyses found significant genetic diversity among regions (69%) and among catchments (58%). Most population pairwise F_ST tests were significant, with non‐significant pairwise tests among sites in the central region and pairs of sites between neighbouring streams. 4. The levels of sequence divergence are interpreted as the result of Pleistocene divergence in multiple refugia, leading to the evolution of regionally unique haplotypes. The low diversity in some central region populations may result from recent colonisation following local extinctions, associated with volcanic events.