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1.
Mitochondrial DNA (mtDNA) evolution was investigated in skinks of the genus Chalcides found in the Canary Islands ( Ch. sexlineatus, Ch. viridanus and Ch. simonyi ), together with some North African congenerics ( Ch. polylepis and Ch. mionecton ). Several sites were included within islands to cover areas of known within-island geographical variation in morphology. Skinks from the islands of El Hierro and La Gomera appear to be sister taxa. The relationships between this clade and the Tenerife and Gran Canarian skinks were not fully resolved, although the best working hypothesis indicated monophyly with the former, with the latter forming a closely related outgroup. Ch. simonyi from Fuerteventura was more distantly related to the Western Canary Island skinks and did not show close relationships with the North African species Ch. mionecton and Ch . polylepis . Possible colonization sequences for the four most Western Canary Islands were considered. El Hierro appears to have been colonized relatively recently from La Gomera, commensurate with the recent origin of this island, while dispersal between La Gomera and Tenerife and between Gran Canaria and Tenerife or La Gomera appears to have taken place considerably earlier. Substantial within-island haplotype divergence was found in Gran Canaria and Tenerife. This may be a result of recent periods of intense volcanic activity found within these two islands. Lower levels of within-island differentiation are found in La Gomera and El Hierro and may be explained by lower levels of volcanic activity during recent geological history and a more recent colonization, respectively.  相似文献   

2.
The processes of island colonization and speciation are investigated through mtDNA studies on Canary Island beetles. The genus Nesotes (Coleoptera: Tenebrionidae) is represented by 19 endemic species on the Canary Islands, the majority of which are single island endemics. Nesotes conformis is the most widespread, occurring on Gran Canaria, Tenerife, La Palma and El Hierro. Nesotes conformis forms a paraphyletic assemblage, with a split between Gran Canaria and the other three islands. Nesotes conformis of the western Canary Islands cluster with Nesotes altivagans and Nesotes elliptipennis from Tenerife. Fifty‐two individuals from this western islands species complex have been sequenced for 675 base pairs of the mtDNA cytochrome oxidase II gene, representing Tenerife, La Palma and El Hierro. A neighbour joining analysis of maximum likelihood distances resulted in three distinct mtDNA lineages for N. conformis, two of which also include mitotypes of N. altivagans and N. elliptipennis. Through application of parametric bootstrap tests, we are able to reject hypotheses of monophyly for both N. conformis and N. altivagans. Nesotes altivagans and N. elliptipennis are poorly separated morphologically and mtDNA sequence data adds support to this being one species with a highly variable morphology. We propose that N. altivagans/N. elliptipennis is recently derived from two ancestral mtDNA lineages within N. conformis from the Teno region of Tenerife. We further propose colonization of the younger islands of La Palma and El Hierro by N. conformis from a mitochondrial lineage within the Teno massif (colonization; diversification; mitochondrial DNA; Canary Islands; Coleoptera).  相似文献   

3.
Chamaecytisus proliferus (L.fil.) Link (Fabaceae: Genisteae) represents a species complex in the Canary Islands. Floristic data from 147 releves from the whole complex were collected and analysed by classification (TWINSPAN) and ordination (DECORANA) methods. Results indicate that white escobon of Tenerife, escobon of El Hierro, white escobon of Gran Canaria and typical tagasaste in La Palma are associated with those plant communities from the north of these islands which are under the influence of the north-eastern trade winds. Narrow-leaved escobon in Tenerife and La Gomera, escobon of southern Gran Canaria and white tagasaste of La Palma are found in those areas which are not under the direct influence of these winds. Morphological forms from the more easterly islands (Gran Canaria and Tenerife-La Gomera) have the broadest ecological range and they have played an important role in the floristic changes which have taken place after the destruction of the forests in these islands. The highest priorities for in situ conservation should be given to wild populations of typical tagasaste, white escobon of Tenerife and escobon of El Hierro.Abbreviations IBPGR International Board for Plant Genetic Resources - DECORANA Detrended Correspondence Analysis - OTU Operational Taxonomic Unit - TWINSPAN Two Way Indicator Species Analysis  相似文献   

4.
Aim The aim of this paper is to identify the patterns in the morphological differentiation in Canary Island mice, based on fossil and modern samples. In order to achieve this, the mouse species present on the archipelago were first compared with a set of continental mice. The differences between the continental and Canary Island samples, and among the Canary Island samples, provide insights into the processes of colonization and the subsequent insular evolution. Location Canary archipelago. Methods An outline analysis based on Fourier transformation was used to quantify shape differences between lower molars. Together with the fossil and modern Canary Island samples, a reference set of genotyped continental populations of the commensal Mus musculus and the wild Mus spretus was used for comparison. Results The morphometric analysis showed that all the mouse specimens from the Canary Islands and Cape Verde belonged to Mus musculus domesticus. Lower molars of extant mice from La Gomera, El Hierro, Gran Canaria, Tenerife, and to a lesser degree from Lanzarote, were similar to those of genotyped M. m. domesticus from the continent, while teeth of extant mice from Fuerteventura were more divergent. Fossil mice from Fuerteventura were very similar to the extant representatives on this island, and similar to the fossil mice on the nearby islands of Lobos and La Graciosa. Main conclusions The mouse present on the Canary archipelago has been identified as the house mouse M. m. domesticus. Based on the shape of the lower molar, the Canary Island mice are divergent from the continental ones, but the degree of divergence varies with the geography of the archipelago. Overall, populations from eastern islands are more divergent from the continental mice than populations from western ones. Fossil populations indicate that this situation was established several centuries ago. Two main factors may have contributed to this pattern: the appearance of different types of environment on the islands since the successful settlement of the mouse, and/or the number of subsequent introductions of continental individuals via shipping.  相似文献   

5.
Many studies have addressed evolution and phylogeography of plant taxa in oceanic islands, but have primarily focused on endemics because of the assumption that in widespread taxa the absence of morphological differentiation between island and mainland populations is due to recent colonization. In this paper, we studied the phylogeography of Scrophularia arguta, a widespread annual species, in an attempt to determine the number and spatiotemporal origins of dispersal events to Canary Islands. Four different regions, ITS and ETS from nDNA and psbA‐trnH and psbJ‐petA from cpDNA, were used to date divergence events within S. arguta lineages and determine the phylogenetic relationships among populations. A haplotype network was obtained to elucidate the phylogenetic relationships among haplotypes. Our results support an ancient origin of S. arguta (Miocene) with expansion and genetic differentiation in the Pliocene coinciding with the aridification of northern Africa and the formation of the Mediterranean climate. Indeed, results indicate for Canary Islands three different events of colonization, including two ancient events that probably happened in the Pliocene and have originated the genetically most divergent populations into this species and, interestingly, a recent third event of colonization of Gran Canaria from mainland instead from the closest islands (Tenerife or Fuerteventura). In spite of the great genetic divergence among populations, it has not implied any morphological variation. Our work highlights the importance of nonendemic species to the genetic richness and conservation of island flora and the significance of the island populations of widespread taxa in the global biodiversity.  相似文献   

6.
FRANCISCO-ORTEGA, J., JACKSON, M. T., SANTOS-GUERRA, A. & FORD-LLOYD, B. V., 1993. Morphological variation in the Chamaecytisus proliferus (L.f.) Link complex (Fabaceae: Genisteae) in the Canary Islands . A multivariate study (Principal Component Analysis and Cluster Analysis, Warďs method) of 47 morphological traits from 164 populations of Chamaecytisus proliferus (L.f.) Link from the Canary Islands confirmed that this species complex is formed by seven morphological types. At least eight traits discriminated between these types. Patterns of variation follow a cline within Gran Canaria, Tenerife and La Palma. These results also show that morphological variation is greater in the eastern islands (i.e. Gran Canaria and Tenerife) than in the western islands (La Gomera, El Hierro and La Palma) and that no morphological differences are found between plants of typical tagasaste from wild and cultivated populations.  相似文献   

7.
The widespread common chaffinch (Fringilla coelebs) inhabits five of the seven Canary Islands. Sequences of the mitochondrial cytochrome b gene (1002 bp) revealed new insights into the systematics and phylogeography of this taxon. Additionally, a set of microsatellite loci were analyzed to examine the structure of these populations. Our results suggest that a new species of the genus Fringilla is present in the Canary Islands, which comprises at least three subspecies, but with a different distribution to that which has been morphologically accepted. The specimens from Gran Canaria are genetically distinct from those of La Gomera and Tenerife (F. c. canariensis), which suggests the existence of an undescribed taxon. Furthermore, nuclear microsatellite data suggest an ongoing incipient speciation process in this population. This study provides both important conservationist implications and a basis for re-evaluating the taxonomic status of the Canarian Fringilla coelebs populations.  相似文献   

8.
Abstract.— The genus Brachyderes Schönherr (Coleoptera: Curculionidae) is represented by the species B. rugatus Wollaston on the Canary Islands, with one subspecies on each of the islands of Gran Canaria, Tenerife, La Palma, and El Hierro. These four subspecies are associated with the endemic pine tree Pinus canariensis , and their distributions are broadly coincident. Eighty-eight individual Canarian Brachyderes , sampled from across the distributions of each subspecies, have been sequenced for 570 bp of the mitochondrial DNA (mtDNA) cytochrome oxidase II gene (COII). No mitotypes are shared among islands. Both maximum-likelihood and distance-based phylogenetic analyses have shown that: Tenerife is composed of a single monophyletic clade of mitotypes, El Hierro is composed of a single monophyletic clade occurring within a larger clade comprising all the La Palma mitotypes, and the mitotypes of these three islands form a monophyletic group distinct from Gran Canaria. New methods for estimating divergence times without the assumption of rate constancy have been used to reconstruct the direction and approximate timing of colonizations among the islands. Colonization has occurred from older to progressionally younger islands, and these colonizations are estimated to have occurred less than 2.6 million years ago, although the timing of the initial colonization of the archipelago is not discernable. New methods for the estimation of diversification rates that use branching times as the analyzed variable have been applied to each island fauna. Hypothesized effects of different levels of recent volcanism among islands were not apparent. All islands exhibit a gradually decreasing rate of genetic diversification that is marked by periodic sudden changes in rate.  相似文献   

9.
An analysis of the sequences of the mitochondrial cytochrome b gene (1005 bp) of the Parus teneriffae-group from the Canary Islands and North Africa revealed new insights into the phylogeography of this taxon. The origin of the radiation on the Canarian Archipelago was apparently one of the central islands—Tenerife or Gran Canaria. The populations on El Hierro (P. t. ombriosus) and La Palma (P. t. palmensis) represent distinct monophyletic lineages. Blue tits from Gran Canaria are genetically distinct from those of La Gomera and Tenerife (P. t. teneriffae), which supports the results of other studies and suggests the existence of an—until now—undescribed taxon there. In contrast, the populations on the eastern islands of Fuerteventura and Lanzarote (P. t. degener) could not be distinguished from North African blue tits (P. t. ultramarinus), and these populations should be subsumed under the subspecies ultramarinus. Taxonomic recommendations based on these results include the distinction of the northern European P. caeruleus from P. teneriffae, including blue tits from North Africa and the Canary Islands, the treatment of degener and ultramarinus as synonymous (P. teneriffae ultramarinus) and a new blue tit taxon on the island of Gran Canaria (P. t. hedwigii nov. ssp.), which is formally described. The genetic results are in parts supported by bioacoustic and morphological data.  相似文献   

10.
The mite genus Steganacarus is represented in the Canary Islands by three endemic species, one recently discovered species, and several morphotypes of uncertain taxonomic position. We used a fragment of the mitochondrial cytochrome oxidase I gene to reconstruct the phylogenetic relationships among representatives of the different taxa from the three central islands of the archipelago, Tenerife, La Gomera and Gran Canaria. Sequence data were analysed by both maximum parsimony and maximum likelihood methods. The inferred phylogenetic relationships do not correlate well with current morphological taxonomy but reveal four deeply divergent and geographically coherent lineages, one each on Gran Canaria and La Gomera and two on Tenerife. No pattern of molecular differentiation was observed among different morphotypes. Possible explanations for this incongruence are suggested in relation to the ecology and biogeography of the group. A recently discovered Steganacarus species from La Gomera, morphologically quite distinct from the other Canarian Steganacarus, is clearly identified as a taxon distantly related to all the other Canarian samples.  相似文献   

11.
Abstract.  We investigated the phylogenetic patterns, evolutionary processes, and their taxonomic implications, of two closely related shield-backed katydid genera endemic to the Macaronesian archipelagos: the monotypic Psalmatophanes Chopard, 1938 endemic to Madeira and Calliphona Krauss, 1892, which includes three species restricted to the Canary Islands. Two main hypotheses have been proposed to explain the origin and colonization pathways of these two genera: a single origin with subsequent sequential colonization of the islands, or three independent colonization waves from continental Africa. We used DNA sequence information from the mitochondrial genes cox1, tRNAleucine, rrnL and nad1 to infer phylogenetic relationships among Psalmatophanes and Calliphona species. Our results provide support for the independent colonization of Madeira and the Canary Islands, and suggest that Psalmatophanes is actually more closely related to the continental genus Tettigonia than to the Canarian representatives. Deep genetic divergence among Canarian species provides further support for the assignment of the Canarian species into two subgenera. Tree topology along with Bayesian-based estimates of lineage age suggest a pattern of colonization from Tenerife to La Palma, and from Tenerife to Gran Canaria with subsequent dispersal to La Gomera. We report the first collection of a Calliphona specimen in the island of El Hierro, which molecular data suggest is a recent immigrant from La Gomera. We hypothesize that the patterns of distribution and genetic divergence exhibited by Calliphona in the Canary Islands are compatible with a taxon cycle process. Our results have further implications for the higher level phylogeny of the subfamily Tettigoniinae and suggest that some of the tribes as currently delimited may not correspond to natural groups.  相似文献   

12.
The Canary Island pine weevil Brachyderes rugatus (Wollaston) consists of four allopatric subspecies that are thought to have arisen from several historic colonization events within the archipelago. We have isolated and optimized seven microsatellite loci from Brachyderes rugatus calvus from Gran Canaria. Six of these loci are polymorphic within B. rugatus (11–22 alleles per locus; heterozygosity between 0.43 and 0.84). There is no evidence for heterozygote deficit within populations or for linkage disequilibrium between pairs of loci. These molecular markers are likely to prove useful tools for quantifying the genetic variability of bottlenecked island populations.  相似文献   

13.
The phylogeny and population history of Meladema diving beetles (Coleoptera, Dytiscidae) were examined using mitochondrial DNA sequence from 16S ribosomal RNA and cytochrome oxidase I genes in 51 individuals from 22 populations of the three extant species (M. imbricata endemic to the western Canary Islands, M. lanio endemic to Madeira and M. coriacea widespread in the Western Mediterranean and on the western Canaries), using a combination of phylogenetic and nested clade analyses. Four main lineages are observed within Meladema, representing the three recognized species plus Corsican populations of M. coriacea. Phylogenetic analyses demonstrate the sister relationship of the two Atlantic Island taxa, and suggest the possible paraphyly of M. coriacea. A molecular clock approach reveals that speciation within the genus occurred in the Early Pleistocene, indicating that the Atlantic Island endemics are not Tertiary relict taxa as had been proposed previously. Our results point to past population bottlenecks in all four lineages, with recent (Late-Middle Pleistocene) range expansion in non-Corsican M. coriacea and M. imbricata. Within the Canary Islands, M. imbricata seems to have independently colonized La Gomera and La Palma from Tenerife (although a colonization of La Palma from La Gomera cannot be discarded), and M. coriacea has independently colonized Tenerife and Gran Canaria from separate mainland lineages. In the Mediterranean basin this species apparently colonized Corsica on a single occasion, relatively early in its evolutionary history (Early Pleistocene), and has colonized Mallorca recently on multiple occasions. On the only island where M. coriacea and M. imbricata are broadly sympatric (Tenerife), we report evidence of bidirectional hybridization between the two species.  相似文献   

14.
Phylogenetic relationships in the Olea europaea complex and the phylogeography of 24 populations of the Macaronesian olive (O. europaea ssp. cerasiformis) were assessed by using three molecular markers: nuclear ribosomal internal transcribed spacer 1 (ITS-1) sequences, randomly amplified polymorphic DNAs (RAPD), and intersimple sequence repeats (ISSR). Parsimony analysis of the ITS-1 sequences and Neighbour-joining (NJ) analyses of RAPD and ISSR banding variation revealed four major lineages in the O. europaea complex: (1) ssp. cuspidata; (2) ssp. cerasiformis from Madeira; (3) ssp. laperrinei; and (4) ssp. cerasiformis from the Canary Islands plus ssp. europaea. These results provide unequivocal support for two independent dispersal events of Olea to the Madeira and Canary Islands. Molecular and morphological evidence led to recognition of two separate olive taxa in Macaronesia, to date included in ssp. cerasiformis. NJ analyses of the combined RAPD and ISSR data suggest that the colonization of the Canaries by O. europaea may have followed an east to west stepping-stone model. An interisland dispersal sequence can be recognized, starting from the continent to Fuerteventura, Gran Canaria, Tenerife, La Gomera, and finally La Palma. High dispersal activity of the lipid-rich Olea fruits by birds in the Mediterranean region is congruent with multiple dispersal of olives to Macaronesia and successive colonization of the archipelagos. The observation of strong genetic isolation between populations of different islands of the Canary Islands suggests, however, that subsequent interisland dispersal and establishment has been very rare or may not have occurred at all.  相似文献   

15.
Geographical isolation by oceanic barriers and climatic stability has been postulated as some of the main factors driving diversification within volcanic archipelagos. However, few studies have focused on the effect that catastrophic volcanic events have had on patterns of within‐island differentiation in geological time. This study employed data from the chloroplast (cpDNA haplotypes) and the nuclear (AFLPs) genomes to examine the patterns of genetic variation in Canarina canariensis, an iconic plant species associated with the endemic laurel forest of the Canary Islands. We found a strong geographical population structure, with a first divergence around 0.8 Ma that has Tenerife as its central axis and divides Canarian populations into eastern and western clades. Genetic diversity was greatest in the geologically stable ‘palaeo‐islands’ of Anaga, Teno and Roque del Conde; these areas were also inferred as the ancestral location of migrant alleles towards other disturbed areas within Tenerife or the nearby islands using a Bayesian approach to phylogeographical clustering. Oceanic barriers, in contrast, appear to have played a lesser role in structuring genetic variation, with intra‐island levels of genetic diversity larger than those between‐islands. We argue that volcanic eruptions and landslides after the merging of the palaeo‐islands 3.5 Ma played key roles in generating genetic boundaries within Tenerife, with the palaeo‐islands acting as refugia against extinction, and as cradles and sources of genetic diversity to other areas within the archipelago.  相似文献   

16.
The Stone-curlew Burhinus oedicnemus, a steppe bird species, is mainly distributed in the Mediterranean and Macaronesian regions, which are considered hotspots of biodiversity with priority for animal and plant species richness conservation. In this study, we investigated the genetic diversity of the Stone-curlew in the Mediterranean basin and in the Canary Islands by applying a multilocus approach. We analysed mitochondrial and nuclear markers in order to evaluate the genetic structure and the congruence between morphological subspecies and geographic samples. We found a significant level of genetic differentiation between Mediterranean and Canary Island populations with all markers. Both in the Mediterranean basin and in the Canary Islands, we found a significant level of genetic diversity with nuclear markers only. We identified seven population groups, including insular populations. The four subspecies described for the Western Palaearctic were confirmed with some changes in distribution range. In spite of habitat fragmentation and negative population trend, the Stone-curlew showed a significant level of genetic diversity and gene flow among continental populations. However, islands constitute important reservoirs of genetic diversity and a potential for the evolution of the species.  相似文献   

17.
Random amplified polymorphic DNA (RAPD) marker variation was analyzed in female gametophytes in natural populations of Gelidium canariense (Grunow) Seoane‐Camba ex Haroun, Gil‐Rodríguez, Diaz de Castro et Prud'Homme van Reine from the Canary Islands to estimate the degree and distribution of genetic variability and differentiation. A total of 190 haploid individuals were analyzed with 60 polymorphic RAPDs bands which produced 190 distinct multilocus genotypes. A high level of polymorphism was detected in all populations analyzed. Within‐population gene diversity ranged from 0.156 to 0.264. The populations on the island of Gran Canaria showed higher genetic variation than the other populations analyzed. The partitioning of molecular variance by analysis of molecular variance showed that most genetic variation resides within populations (68.85%). These results suggest that sexual reproduction is the predominant mode of reproduction for G. canariense gametophytic populations, and the main determinant in reaching high levels of genetic diversity. The Neighbor‐Joining tree and FCA analysis displayed two subclusters that correspond to the populations from the western islands (Tenerife, La Palma, Gomera) and the eastern island (Gran Canaria). In addition, we have detected a significant relationship between FST/(1?FST) and geographical distance consistent with data on water circulation and age of islands. The results obtained agree with an isolation by distance model, with gene flow from eastern to the western islands, and a high level of genetic differentiation between populations (FST=0.311, P<0.001).  相似文献   

18.
This contribution aimed to predict the invasive Barbary ground squirrel (Atlantoxerus getulus) potentiality for invading the Canary Islands and western Mediterranean region, by determining firstly the climatic suitable areas in its native range and secondly, using presence data in the invaded range. Nineteen environmental variables submitted to a Principal Components Analysis selected those variables with higher factor loadings, which represent the main environmental conditions of the Northern African region (temperature in the coldest quarter, seasonal temperature, precipitation in the coldest quarter, temperature in the wettest quarter). After selecting hundred times more pseudo-absence points than presence observations (n = 6600 at a 0.083° resolution), Generalized Additive Models and Single-hidden-layer Neural Networks fitted in R were used to calibrate the model. Model results were extrapolated for the Canary Islands and the western Mediterranean region. In order to select between the two techniques, we calculated three accuracy measures (specificity, sensitivity and AUC) after using a Jack-knifing procedure and models were repeated ten times. The GAM model was less accurate than the NN model. Suitable areas did not have mean temperatures in the coldest quarter lower than −5°C and precipitation in the coldest quarter higher than 300 mm, respectively. We predicted favorable climatic areas across almost all the Maghreb, the European western Mediterranean region and in all the Canary Islands. Nevertheless, the seven islands differed significantly in the mean favorability scores, with El Hierro, Lanzarote and Gran Canaria being the most suitable. Same methodological analysis was applied to predict A. getulus distribution in other Canarian islands based on presence data from the invaded Fuerteventura. In this case, only Lanzarote and Gran Canaria appeared to be climatically suitable for the species. Our predictive model is an applicable tool to establish the invasive potential of A. getulus and to prioritize management strategies, within and outside the Canarian archipelago, to impede the expansion of this invasive squirrel out of Fuerteventura Island.  相似文献   

19.
Sequences from fragments of the 12S ribosomal RNA and cytochrome b mitochondrial genes were used to analyze phylogenetic relationships among geckos of genus Tarentola from the Canary Islands. A surprisingly high level of within island differentiation was found in T. delalandii in Tenerife and T. boettgeri in Gran Canaria. Molecular differentiation between populations of T. angustimentalis on Lanzarote and Fuerteventura, and between Moroccan and Iberian Peninsula T. mauritanica, also indicate that at least two subspecies should be recognized within each of them. Phylogenetic relationships among these species reveals a higher level of differentiation and a more complex colonization pattern than those found for the endemic genus Gallotia. Lack of evidence for the presence of T. boettgeri bischoffi on the island of Madeira does not seem to support the origin of T. delalandii, T. gomerensis and the canarian subspecies of T. boettgeri from this island, whereas molecular data confirms that T. angustimentalis is a sister species of the continental T. mauritanica. Several independent colonization events from the continent and the extinction of some species are probably responsible for the current distribution of Tarentola in the Canary Islands.  相似文献   

20.
Oceanic archipelagos of volcanic origin have been important in the study of evolution because they provide repeated natural experiments allowing rigorous tests of evolutionary hypotheses. Ongoing volcanism on these islands may, however, affect the evolutionary diversification of species. Analysis of population structure and phylogeographic patterns in island populations can provide insight into evolutionary dynamics on volcanic islands. We analysed genetic and morphological variation in the gecko Tarentola boettgeri on the island of Gran Canaria and compared it with Tarentola delalandii on Tenerife, a neighbouring volcanic island of similar age but distinctly different geological past. Intraspecific divergence of mitochondrial haplotypes indicates long-term persistence of Tarentola on each island, with a phylogeographic signal left by older volcanic events. More recent volcanic eruptions (approximately 0.2 million years ago on Tenerife, approximately 2.2 million years ago on Gran Canaria) have left a signature of population expansion in the population genetic structure, the strength of which depends on the time since the last major volcanic eruption on each island. While these stochastic events have left traces in morphological variation in Tenerife, in Gran Canaria geographical variation was solely associated with environmental variables. This suggests that historically caused patterns in morphology may be overwritten by natural selection within 2 million years.  相似文献   

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