Phylogeny and biogeography of the Petaurista philippensis complex (Rodentia: Sciuridae), inter- and intraspecific relationships inferred from molecular and morphometric analysis

With modified DNA extraction and purification protocols, the complete cytochrome b gene sequences (1140 bp) were determined from degraded museum specimens. Molecular analysis and morphological examination of cranial characteristics of the giant flying squirrels of Petaurista philippensis complex (P. grandis, P. hainana, and P. yunanensis) and other Petaurista species yielded new insights into long-standing controversies in the Petaurista systematics. Patterns of genetic variations and morphological differences observed in this study indicate that P. hainana, P. albiventer, and P. yunanensis can be recognized as distinct species, and P. grandis and P. petaurista are conspecific populations. Phylogenetic relationships reconstructed by using parsimony, likelihood, and Bayesian methods reveal that, with P. leucogenys as the basal branch, all Petaurista groups formed two distinct clades. Petaurista philippensis, P. hainana, P. yunanensis, and P. albiventer are clustered in the same clade, while P. grandis shows a close relationship to P. petaurista. Deduced divergence times based on Bayesian analysis and the transversional substitution at the third codon suggest that the retreating of glaciers and upheavals or movements of tectonic plates in the Pliocene-Pleistocene were the major factors responsible for the present geographical distributions of Petaurista groups.     

基于cpDNA trnL-F序列的胡黄连保护遗传学研究

胡黄连为特产中国-喜马拉雅特有高山植物,作为常用中、藏药材,受到灭绝性采挖,作为濒危和二级保护植物亟待科学的保护。该研究以云南和西藏7个野生居群91个个体为材料,基于 cpDNA trnL-F 非编码序列测序分析胡黄连的遗传多样性和遗传结构,分析显著进化单元,确立优先保护居群并提出科学的保护策略。结果表明：胡黄连 trnL-F 序列长度为871~876 bp,根据序列的核苷酸变异共鉴定出5个单倍型,西藏占有2个单倍型,云南占有3个单倍型,西藏和云南2个地区的所有单倍型均不共享。胡黄连具有较低的单倍型多样性(Hd ＝0.43419)和核苷酸多样性(Dij ＝0.00466)。种群间分化度(Fst ＝0.864520)和基因流(Nm ＝0.04)、居群间的遗传分化水平(GST ＝0.916)、AMOVA 分析(0.78％的遗传变异发生在居群内,60.97％的遗传变异发生在地区内居群间,38.25％的遗传变异发生在地区间)均表明,胡黄连居群间存在明显遗传分化。多数一致性树将胡黄连划分为3个进化分支(Ⅰ、Ⅱ、Ⅲ),这3个分支均与地理相关,分支Ⅰ分布于横断山区的4个居群,分支Ⅱ是分布于东喜马拉雅的一个居群,分支Ⅲ是分布于喜马拉雅中段的2个居群。3个分支分属于3个“进化显著单元(ESU)”。这3个 ESU 中白马雪山、茨中、定日、波密、聂拉木五个居群都需要保护,建议现阶段应优先保护的居群是云南白马雪山和西藏波密居群,以就地保护为主。     

Isozyme variation inFaidherbia albida (Leguminosae, Mimosoideae)

Genetic variation has been assessed in 30 populations (931 families) ofFaidherbia albida (Leguminosae, Mimosoideae) from across its entire African range, using six isozyme loci identified by five enzyme systems. Among the populations studied a null allele was proposed to explain the absence ofLap-1 activity in populations from southern and eastern Africa. The mean percentage of polymorphic loci per population, the mean number of alleles per locus and the mean genetic diversity within populations were 31.7%, 1.6 and 0.127 respectively. Genetic diversity was greatest in populations from West Africa and lowest in populations from eastern/southern Africa, with Ethiopian/Sudanese populations intermediate. The overall degree of genetic differentiation between populations (G_ST) indicated that approximately 56% of the enzyme variation resided within populations. Clustering of Nei's unbaised genetic distances calculated between all populations produced a dendrogram that generally followed the geographic distribution of the populations. Two major groups were identified that may be considered the eastern/southern African and the Ethiopian/West African clusters. Within the Ethiopian/West African cluster two subclusters could be recognised, one broadly corresponding to those populations from Ethiopia/Sudan and the other to those populations from West Africa. The implications of these results for theories regarding the origin ofF. albida in Africa are discussed.     

The contact zone between two migration routes of silver fir,Abies alba (Pinaceae), revealed by allozyme studies

Enzyme data from 35 populations or regional samples ofAbies alba were obtained from different regions of Austria and Bavaria. The genetic analysis revealed a clear differentiation between the western and the north/eastern populations. A rather narrow transition zone (the river valleys of Salzach and Traun) with genetically intermediate populations connects these two regions. This west-east differentiation can be interpreted as the result of distinct migration routes from a glacial refuge in Central Italy. The transition zone is assumed to be the contact area of two routes north of the Alps. Each migration route of silver fir parallels the distribution range ofCardamine trifolia (east) andAposeris foetida (west) respectively. It is supposed that these two species were already associated with silver fir during colonization.Dedicated to emer. Univ.-Prof. DrFriedrich Ehrendorfer on the occasion of his 70th birthday     

Connectivity of populations within and between major biogeographic regions of the tropical Pacific in <Emphasis Type="Italic">Conus ebraeus</Emphasis>, a widespread marine gastropod

Information on genetic connectivity and structure of populations in the tropical Pacific is critical for making inferences about the origins and maintenance of diversity in this region. Sequences of the mitochondrial COI gene from 92 individuals of the trans-Pacific gastropod Conus ebraeus from eight localities spanning the tropical Pacific were analyzed to determine whether populations in the western, central, and eastern Pacific exhibit genetic structure, to examine the demographic histories of populations, and to infer patterns of gene flow. A total of 43 unique haplotypes were recovered, including a common haplotype that occurred in six of the eight populations examined. AMOVA and pairwise F-statistics showed that populations in the western and central Pacific were significantly differentiated from populations in the eastern Pacific, but no other evidence of structure. Bayesian isolation–migration (IM) analysis suggested that populations in the western and central Pacific separated from those in eastern Pacific during the Pleistocene. Examination of mismatch distributions and results from IM revealed that populations in the western and central Pacific expanded during the Pleistocene. Gene flow across the East Pacific Barrier appears to occur predominantly westward.     

Genetic variability and the effect of habitat fragmentation in spotted suslik Spermophilus suslicus populations from two different regions

Endangered species worldwide exist in remnant populations, often within fragmented landscapes. Although assessment of genetic diversity in fragmented habitats is very important for conservation purposes, it is usually impossible to evaluate the amount of diversity that has actually been lost. Here, we compared population structure and levels of genetic diversity within populations of spotted suslik Spermophilus suslicus, inhabiting two different parts of the species range characterized by different levels of habitat connectivity. We used microsatellites to analyze 10 critically endangered populations located at the western part of the range, where suslik habitat have been severely devastated due to agriculture industrialization. Their genetic composition was compared with four populations from the eastern part of the range where the species still occupies habitat with reasonable levels of connectivity. In the western region, we detected extreme population structure (F _ST = 0.20) and levels of genetic diversity (Allelic richness ranged from 1.45 to 3.07) characteristic for highly endangered populations. Alternatively, in the eastern region we found significantly higher allelic richness (from 5.09 to 5.81) and insignificant population structure (F _ST = 0.03). As we identified a strong correlation between genetic and geographic distance and a lack of private alleles in the western region, we conclude that extreme population structure and lower genetic diversity is due to recent habitat loss. Results from this study provide guidelines for conservation and management of this highly endangered species.     

Phylogenetic relationships of the Japanese minnows,Pseudorasbora (Cyprinidae), as inferred from mitochondrial 16S rRNA gene sequences

The phylogenetic relationships among threePseudorasbora fishes (Cyprinidae, Sarcocheiichthyinae) occurring in Japan (P. parva, P. pumila pumila andP. pumila subsp. sensu Nakamura [1963]) were inferred from nucleotide sequences of the mitochondrial 16S rRNA gene. The sequences. of 1240 bp, were determined and compared for 22 specimens from 2–8 populations for each taxon, with a singlePungtungia herzi specimen as an outgroup. A total of 171 sites (13.8%) were variable among the specimens, but only 0–2 sites within each population. The phylogenetic relationships estimated by neighbor-joining, maximum-parsimony and maximum-likelihood methods confirmed a sister relationship between the twoP. pumila subspecies, with a high level of confidence. However, their genetic distinction from each other (4.1±0.4SD % sequence difference on average) was at a level similar to that between them andP. parva (5.9±0.5%). The geographic distribution of the twoP. pumila subspecies, which are separated by the Fossa Magna region, suggests that the genetic divergence of the two subspecies originated from a vicariant process separating the freshwater ichthyofaunas of eastern and western Honshu.Pseudorasbora parva populations were divided into two genetic groups (1.8±0.2% sequence difference), one group comprising continental and part of the Japanese populations, and the other the remaining Japanese populations. This suggests that at least two genetically divergent lineages had been originally distributed in Japan, but a strong possibility remains that the present situation has resulted from artificial transplantation.     

Population structure of the sailfin sandfish, Arctoscopus japonicus (Trichodontidae), in the Sea of Japan

We analyzed patterns of genetic diversity in the sailfin sandfish (Arctoscopus japonicus), focusing on population subdivisions within the Sea of Japan. We observed 270 specimens from nine sampling sites in 1999–2000, i.e., seven sites in the Sea of Japan and two sites from the Pacific coast of Hokkaido. An additional site (30 specimens) was sampled from eastern Korea in the spawning season of 2004 for comparison. Forty haplotypes, compiled into three haplogroups (A–C), were detected based on the comparison of a 400-bp sequence of the anterior part of the mitochondrial control region. In accordance with previous hypotheses from morphological and molecular analyses, genetic discontinuity between the Sea of Japan and the Pacific coast of Hokkaido was conspicuous. Within the Sea of Japan, eight sampling sites were not genetically uniform, and most of the variations among sites were detected between eastern Korea [the “eastern Korea” (EK) population: distributed from the Korean Peninsula to Mishima, Yamaguchi Prefecture] and the other sites along the coast of Japan [the “western Japan” (WJ) population: from Oki Islands to western Hokkaido] (Φ _CT  = 0.096, P = 0.0183). The WJ population, having lower genetic variability, showed significant departure from neutrality, indicating influences through a recent population expansion. The period of the expansion can be estimated to have begun on the order of 10⁴ years ago. We consider that the present Japan Sea populations have been formed through the invasion of a small ancestral stock to the Sea of Japan and its population expansion during the last glacial period or later. On the other hand, we failed to detect distinct evidence of a population expansion in the EK population. Haplogroup C, detected in a high frequency in this population, was estimated to have mixed with haplogroup A after rapid differentiations of the latter. Therefore, the EK population, strongly influenced by such a mixture, might possess haplogroup C in a higher frequency and a different haplotype composition from the WJ population.     

Geographical patterns of deep mitochondrial differentiation in widespread Malagasy reptiles

Using sequences of the mitochondrial 16S rRNA gene, we reconstructed the phylogeography of six widely distributed Malagasy reptiles: two gekkonid lizard species, Phelsuma lineata and Hemidactylus mercatorius; two chameleons, the Calumma brevicorne complex, and Furcifer lateralis; and two skinks, Trachylepis gravenhorstii and Trachylepis elegans. Genetic differentiation among major haplotype lineages was high and in some cases indicates or confirms species status of the divergent populations. Maximum uncorrected sequence divergences were between 2.2% and 8.3% within the various species or species complexes. Haplotype lineages were exclusive to geographic regions, except in the commensal H. mercatorius where in three anthropogenic habitats coexistence of haplotype lineages was observed, possibly due to human translocation. The eastward flowing rivers Mangoro and Mananara may represent barriers to gene flow in the case of three species each. Some species sampled from humid eastern and arid western Madagascar showed no differentiation between populations from these two regions; instead the pattern observed was in several cases more concordant with a differentiation along a north-south axis.     

Dynamics of Mycoplasmal Conjunctivitis in the Native and Introduced Range of the Host

In 1994, Mycoplasma gallisepticum, a common bacterial poultry pathogen, caused an epidemic in house finches in the eastern part of their North American range where the species had been introduced in the 1940s. Birds with mycoplasmal conjunctivitis were reported across the entire eastern United States within 3–4 years. Here we track the course of the Mycoplasma gallisepticum epidemic as it reached native, western North American populations of the house finch. In 2002, Mycoplasma gallisepticum was first observed in a native house finch population in Missoula, MT, where it gradually increased in prevalence during the next 2 years. Concurrently, house finches with conjunctivitis were reported with increasing number in the Pacific Northwest. In native populations of the host, the epidemic expanded more slowly, and reached lower levels of prevalence than in the eastern, introduced range of the species. Maximal prevalence was about half in the Missoula population than in local populations in the East. Although many factors can contribute to these differences, we argue that it is most likely the higher genetic heterogeneity in western than in eastern populations caused the lower impact of the pathogen.     

Conservation genetics of the threatened horned grebe (<Emphasis Type="Italic">Podiceps auritus</Emphasis> L.) population of the Magdalen Islands,Québec

The horned grebe (Podiceps auritus) population of the Magdalen Islands in the St. Lawrence Gulf (Québec, Canada) has declined sharply over the last decades. It is the only breeding population of this species in eastern North America with nearest breeding populations being >2500 km apart in western North America and Europe, We used three types of genetic markers: mitochondrial (mt) DNA ND2 sequence, α-enolase intron sequence, and 25 amplified fragment length polymorphism loci (AFLPs) to quantify the genetic diversity within the Magdalen Island population and to assess its genetic distinctiveness relative to populations from western Canada (five sites) and Iceland (one site). The Magdalen Island population retained a comparable amount of genetic diversity to the average diversity observed across all populations in all three markers. Horned grebe mtDNA sequences formed a monophyletic group and nearly all haplotypes present in Québec were found elsewhere. In the ND2 fragment, populations partitioned into two groups corresponding to subspecies (Iceland versus North American sites) and more strongly in three groups according to geographic disjunctions (Iceland versus Québec versus western Canada). In contrast, there was no evidence of structure between sites in the α-enolase intron. In the AFLPs, Iceland showed the greatest level of differentiation, followed by the Québec and British Columbia populations. For conservation purposes, we suggest that the Magdalen Islands population should be recognized as a separate unit.     

Corresponding patterns of geographic variation among populations ofSilene latifolia (=S. alba =S. pratensis) (Caryophyllaceae)

Morphological and biochemical data were analysed from 30 greenhouse-grown populations of EuropeanSilene latifolia. Six separate character sets (flavones, seed, pollen, capsules, male and female flower morphology) were used in the analyses. There was broad-scale congruence between trends of geographic variation in most character sets, with the populations being assigned to western (or southern and western) and eastern clusters. The eastern and western clusters abut along a transition zone that runs roughly from Belgium to the northern Balkans; this zone represents a region of relatively rapid change and contains populations intermediate between the eastern and western clusters. Variation in flower morphology was weak and discordant with variation in the other character sets. The origin and maintenance of the variation pattern is discussed in terms of migrational history and hybrid zones.     

Range expansion leading to departures from neutrality in the nonsymbiotic hemoglobin gene and the cpDNA trnL-trnF intergenic spacer in Trema dielsiana (Ulmaceae)

The population genetics and phylogeography of Trema dielsiana in Taiwan were inferred from genetic diversity at the nonsymbiotic hemoglobin gene and the trnL-trnF intergenic spacer of cpDNA. Reduced genetic variation was detected in these two unlinked genes. The gene genealogy of the hemoglobin locus recovered two lineages corresponding to the western and eastern regions of Taiwan. This pattern is compatible with a past fragmentation event revealed by phylogeographical analyses. To distinguish between selective departures from neutrality (i.e., heterogeneous processes) and demographic (homogeneous) processes, Hahn et al.'s heterogeneity test was conducted on the hemoglobin gene. Lack of significant differences in Tajima's D statistics between synonymous and nonsynonymous mutations indicates that homogeneous processes may have played a key role in governing the evolution of the functional locus. Significantly negative Tajima's D estimates for both overall exons and introns of the hemoglobin gene as well as for the cpDNA intergenic spacer support a phylogeographical hypothesis of range expansion after genetic bottlenecks. High level of genetic variation and a negative Tajima's D statistic suggests a possible northern refugium that may have harbored populations during the glacial maximum.     

Intraspecific comparison of population structure, genetic diversity, and dispersal among three subspecies of Townsend’s big-eared bats, Corynorhinus townsendii townsendii, C. t. pallescens, and the endangered C. t. virginianus

Townsend’s big-eared bat, Corynorhinus townsendii, is distributed broadly across western North America and in two isolated, endangered populations in central and eastern United States. There are five subspecies of C. townsendii; C. t. pallescens, C. t. australis, C. t. townsendii, C. t. ingens, and C. t. virginianus with varying degrees of concern over the conservation status of each. The aim of this study was to use mitochondrial and microsatellite DNA data to examine genetic diversity, population differentiation, and dispersal of three C. townsendii subspecies. C. t. virginianus is found in isolated populations in the eastern United States and was listed as endangered under the Endangered Species Act in 1979. Concern also exists about declining populations of two western subspecies, C. t. pallescens and C. t. townsendii. Using a comparative approach, estimates of the genetic diversity within populations of the endangered subspecies, C. t. virginianus, were found to be significantly lower than within populations of the two western subspecies. Further, both classes of molecular markers revealed significant differentiation among regional populations of C. t. virginianus with most genetic diversity distributed among populations. Genetic diversity was not significantly different between C. t. townsendii and C. t. pallescens. Some populations of C. t. townsendii are not genetically differentiated from populations of C. t. pallescens in areas of sympatry. For the western subspecies gene flow appears to occur primarily through male dispersal. Finally, geographic regions representing significantly differentiated and genetically unique populations of C. townsendii virginianus are recognized as distinct evolutionary significant units.     

Gene variation and gene flow inOrchis morio (Orchidaceae) from Italy

Data are presented on genetic variation at 27 enzyme loci of the Green-Winged orchid,Orchis morio, in 18 population samples from Italy. The existence in Italy of two subspecies, i.e. subspp.morio andpicta, is not supported by allozyme data. No genetic heterogeneity was found betweenmorio-like andpicta-like samples and specimens. Moreover, morphological transition between the two forms was observed in different Italian populations. The parameters of genetic variability estimated forO. morio populations are consistent with those found among monocotyledon plants, and among those outcrossing, animal-pollinated and with wind-dispersed seeds. Genetic diversity of ItalianO. morio is mostly within populations. Correspondingly, low values of interpopulational genetic distance were found. This appears to be due to high levels of gene flow, which were estimated with different methods. The lack ofO. longicornu from Italian samples, as well as of any hybrid withO. morio (F₁, backcrossed or recombinant individuals) is demonstrated on the basis of genetic data. It is concluded that recurrent reports ofO. longicornu from Italy are due to confusion withO. morio or with otherOrchis species.     

Global population genetic structure of the starlet anemone <Emphasis Type="Italic">Nematostella vectensis</Emphasis>: multiple introductions and implications for conservation policy

Distinguishing natural versus anthropogenic dispersal of organisms is essential for determining the native range of a species and implementing an effective conservation strategy. For cryptogenic species with limited historical records, molecular data can help to identify introductions. Nematostella vectensis is a small, burrowing estuarine sea anemone found in tidally restricted salt marsh pools. This species’ current distribution extends over three coast lines: (i) the Atlantic coast of North America from Nova Scotia to Georgia, (ii) the Pacific coast of North America from Washington to central California, and (iii) the southeast coast of England. The 1996 IUCN Red List designates N. vectensis as “vulnerable” in England. Amplified fragment length polymorphism (AFLP) fingerprinting of 516 individuals from 24 N. vectensis populations throughout its range and mtDNA sequencing of a subsample of these individuals strongly suggest that anthropogenic dispersal has played a significant role in its current distribution. Certain western Atlantic populations of N. vectensis exhibit greater genetic similarity to Pacific populations or English populations than to other western Atlantic populations. At the same time, F-statistics showing high degrees of genetic differentiation between geographically proximate populations support a low likelihood for natural dispersal between salt marshes. Furthermore, the western Atlantic harbors greater genetic diversity than either England or the eastern Pacific. Collectively, these data clearly imply that N. vectensis is native to the Atlantic coast of North America and that populations along the Pacific coast and in England are cases of successful introduction.     

Tyrrhenian basins of Ligury as a new peri-Mediterranean ichthyogeographic district? Population structure of <Emphasis Type="Italic">Telestes muticellus</Emphasis> (Osteichthyes,Cyprinidae), a primary freshwater fish

The phylogeographic structure of vairone (Telestes muticellus), a primary freshwater fish endangered in a large part of its distributional range, was assessed: (i) to reconstruct the complex dispersion pattern in the upper Tyrrhenian hydrographic basins of Ligury, actually not recognised as peri-Mediterranean ichthyogeographic district, and (ii) to evidence the shape of population genetic structure as useful tool for future conservation strategies. A partial fragment of mitochondrial DNA cytochrome b (497 bp) was sequenced in 109 specimens sampled from eight populations, along an east–west geographic gradient. Fourteen haplotypes were identified, confirming the evolutionary distance between the two co-generic species: T. muticellus as ‘Ligurian’ clade and T. souffia as ‘French’ clade. The Nested Clade Analysis (NCA), the population genetic variability and population structure suggested a natural colonization occurred throughout the crossing of Alpine and Apennine watershed. The hierarchical Analysis of Molecular Variance (AMOVA) confirmed a geographic distinction between the populations from western (WTL) and from eastern (ETL) Tyrrhenian basins of Ligury colonised through the river capture processes along the Maritime Alpine watershed (Padano-Venetian district) and along the Apennine watershed (Tuscano-Latium district), respectively. Our results, evidencing the lack of genetic contiguity among vairone populations of the upper Tyrrhenian hydrographic basins of Ligury, allowed to recognise the presence of two management units (MUs) for its conservation. Handling editor: Christian Sturmbauer     

Molecular genetic variation across the southern and eastern geographic ranges of the African lion, <Emphasis Type="Italic">Panthera leo</Emphasis>

We examined sequence variation in the mitochondrial cytochrome b and NADH dehydrogenase subunit 5 genes (2,360 bp total) for 26 lions from eleven locations throughout sub-Saharan Africa. Six distinct haplotypes were observed in the combined sequences, forming two clades: the eastern and the western savannas. The Uganda-Western Kenya haplotype grouped at a basal position with the eastern clade of lions from Tsavo south to the Transvaal and Natal regions. The phylogenetic position of the haplotype from Sabi Sands in the southern part of Kruger National Park remained poorly resolved. The haplotypes found in Namibia and Botswana formed the western clade. The modest genetic variation documented here argues against taxonomic distinctions among living African lions.     

Phylogeography and conservation genetics of Hygrophila pogonocalyx (Acanthaceae) based on atpB–rbcL noncoding spacer cpDNA

Genetic variation in the atpB–rbcL intergenic spacer region of chloroplast DNA (cpDNA) was investigated in Hygrophila pogonocalyx Hayata (Acanthaceae), an endangered and endemic species in Taiwan. In this aquatic species, seed dispersal from capsules via elasticity is constrained by gravity and is thereby confined within populations, resulting in limited gene flow between populations. In this study, a total of 849 bp of the cpDNA atpB–rbcL spacer were sequenced from eight populations of H. pogonocalyx. Nucleotide diversity in the cpDNA is low (=0.00343±0.00041). The distribution of genetic variation among populations agrees with an isolation-by-distance model. Two geographically correlated groups, the western and eastern regions, were identified in a neighbor-joining tree and a minimum-spanning network. Phylogeographical analyses based on the cpDNA network suggest that the present-day differentiation between western and eastern groups of H. pogonocalyx resulted from past fragmentation. The differentiation between eastern and western populations may be ascribed to isolation since the formation of the Central Mountain Range about 5 million years ago, which is consistent with the rate estimates based on a molecular clock of cpDNA.Huang JC and Wang WK equally contributed to this work.     

Differentiation among Austrian populations of Norway spruce [Picea abies (L.) Karst.] assayed by mitochondrial DNA markers

Thirty-seven populations of Norway spruce [Picea abies (L.) Karst.] across the Austrian Alps and Bohemian Massif were sampled to elucidate the geographical pattern of genetic differentiation. Three polymorphic mitochondrial DNA (mtDNA) loci were surveyed. Two or three alleles were detected at each locus, resulting in seven multilocus mtDNA haplotypes (A–F). Western populations proved to be monomorphic, whereas eastern and central Austrian populations were slightly to highly polymorphic, respectively. As revealed by spatial analysis of molecular variance and Monmonier’s analysis, the two main haplotypes A and B are not randomly distributed. Haplotype A was restricted to central and eastern Austria, whereas haplotype B occurred in all Austrian populations but was the only haplotype identified in western populations. This pattern may be explained by different glacial refugia located in the Dinaric Alps and the Carpathian mountains.