Differentiation of subspecies of the polymorphic species Hyalomma marginatum (Acari: Ixodidae) based on immature stages

Study of morphological characters to identify the subspecies of Hyalomma marginatum immature stages is based on material collected throughout all the geographical range of the species. As it was found, the discrimination of subspecies of H. marginatum immature stages needs a complex use of structural and morphometrical characters. Only H. m. marginatum larvae and H. m. isaaci nymphs may be easily differentiated from other subspecies based on the structural (qualitative) characters. Besides, morphometric (quantitative) characters should be used only in combinations for identification of the subspecies. In regard to structural characters of larvae, it was found that the shape and size of coxal spurs of H. marginatum allow differentiating this subspecies from the others (Fig. 3, 5, 6). Based on morphometrical characters of larvae, the following characteristic features of subspecies have been found: in H. m. marginatum, the capitulum is wide, the palpi, hypostome and genua I are short and narrow; in H. m. turanicum, the capitulum is narrow, the palpi are short and narrow, the hypostome is short and wide, the genua I are long and narrow; in H. m. rufipes, the scutum is large, the capitulum is wide, the palpi are long and narrow, the hypostome and genua I are long and wide; in H. m. isaaci, the scutum is small, the capitulum is narrow, the palpi and hypostome are long and narrow, the genua I are short and narrow. Among structural characters of nymph, several discriminative features of subspecies have been found. The shape of the scutum in H. m. isaaci (Fig. 2, 2) clearly differentiates this subspecies from the others (Fig. 2, 1). In H. m. marginatum, the setae of alloscutum as a rule have bluntly rounded apices (Fig. 1, 2), while in the other subspecies these setae are more tapering apically (Fig. 1, 1). The shape of spiracular plates is rather variable within the species, but in H. m. marginatum and H. m. isaaci (Fig. 1, 3) the plates are larger and more perforated than in H. m. rufipes and H. m. turanicum (Fig. 1, 4). In H. m. isaaci, the spurs of coxae I are narrower than in other subspecies (Fig. 1, 8). Based on morphometrical characters of nymphs, the following discriminative features of subspecies have been found: in H. m. marginatum, the capitulum is wide, the palpi are short and wide; in H. m. turanicum, the capitulum is narrow, the palpi are long and narrow; in H. m. rufipes, the scutum is wide, the capitulum is narrow, the palpi are long and narrow, the hypostome is long; in H. m. isaaci, the scutum is long and narrow, the capitulum is wide, the palpi are short and wide, the hypostome is short.     

Larval identification of species and subspecies of the genus Hyalomma (Acari: Ixodidae) from Russia and neighbouring territories

The following species and subspecies of Hyalomma Koch, 1844 are recorded from Russia and neighbouring territories, including those involved in natural foci of tick-borne diseases: H. (Hyalomma) aegyptium (Linnaeus, 1758), H. (Euhyalomma) dromedarii Koch, 1844, H. (Euh.) asiaticum asiaticum Schulze et Schlottke, 1930, H. (Euh.) asiaticum caucasicum Pomerantzev, 1940, H. (Euh.) asiaticum kozlovi Olenev, 1931, H. (Euh.) anatolicum Koch, 1844, H. (Euh.) excavatum Koch, 1844, H. scupense Schulze, 1918, H. (Euh.) marginatum marginatum Koch, 1844, H. (Euh.) marginatum turanicum Pomerantzev, 1946, and H. (Euh.) marginatum rufipes Koch, 1844. The geographic distribution and host-parasite relationships of each taxon are discussed. Species characters of the larval stage, that can be distinguished using light microscopy, are found to be very few. These characters include shape of scutum, shape and rate of hypostome denticulation, shape and rate of the development of spurs on coxae I to III. Measurements of some morphological structure and their rations show statistically significant differences between some closely related species, even if qualitative discriminating characters are unknown. Only measurements and their ratios can be used for the discrimination of larval H. anatolicum from larval H. excavatum, because qualitative discriminating features have not been found for these species. The complex structure of the subspecific morhological differentiation of all parasitic stages in the polymorphic species H. asiaticum and H. marginatum is revealed. This structure probably reflects some peculiarities of the microevolutionary processes. Identification key for the larval stage of seven Hyalomma species is provided.     

Transstadial transmission of Theileria annulata through common ixodid ticks infesting Indian cattle.

Transstadial transmission of Theileria annulata with Hyalomma anatolicum anatolicum, H. dromedarii and H. marginatum isaaci, and Haemaphysalis bispinosa, Rhipicephalus haemaphysaloides haemaphysaloides and Boophilus microplus was determined. It was found that the infection was successfully transmitted by H. a. anatolicum from larva to nymph and nymph to adult in all attempts. When larvae were fed on an infected calf the succeeding adults transmitted the infection when the intervening nymphs fed on a rabbit (non-susceptible host) but not when fed on a calf (susceptible host). Infective adult ticks transmitted the parasite during the first 24 h of feeding on a calf. When the feeding was interrupted after 24 h, and the tick transferred to another calf, the infection was transmitted to the latter as well. H. dromedarii successfully transmitted the infection from larva to nymph and from nymph to adult. Larvae of H. marginatum isaaci did not feed on calves but the infection was successfully transmitted from nymph to adult. Haemaphysalis bispinosa, Rhipicephalus h. haemaphysaloides and Boophilus microplus did not transmit Th. annulata from larva to nymph to adult.     

Molecules and morphology: evidence for cryptic hybridization in African Hyalomma (Acari: Ixodidae)

The role of natural hybridization and introgression as part of the evolutionary process is of increasing interest to zoologists, particularly as more examples of gene exchange among species are identified. We present mitochondrial and nuclear sequence data for Hyalomma dromedarii, Hyalomma truncatum, and Hyalomma marginatum rufipes (Acari: Ixodidae) collected from one-humped camels in Ethiopia. These species are well differentiated morphologically and genetically; sequence data from the mitochondrial DNA (mtDNA) cytochrome oxidase I gene indicates 10-14% divergence between the species. However, incongruence between morphology and the mtDNA phylogeny was observed, with multiple individuals of H. dromedarii and H. truncatum present on the same mtDNA lineage as H. marginatum rufipes. Thus, individuals with morphology of H. dromedarii and H. truncatum are indistinguishable from H. marginatum rufipes on the basis of mtDNA. Multiple copies of ITS-2 were subsequently cloned and sequenced for a subset of individuals from the mtDNA phylogeny, representing both 'normal' and 'putative hybrid' individuals. Very low sequence divergence (0.3%) was observed within 'normal' individuals of both H. dromedarii and H. truncatum relative to the 'putative hybrid' individuals (6 and 2.7%, respectively). The pattern of intra-individual variation in ITS-2 within 'putative hybrid' individuals, particularly in H. dromedarii, strongly suggests that gene flow has occurred among these Hyalomma species, but no indication of this is given by the morphology of the individuals.     

Host-parasite relationships of the genus Hyalomma Koch, 1844 (Acari, Ixodidae) and their connection with microevolutionary process

Host-parasite relationships of Hyalomma species of the world fauna are analyzed. The majority of species infests predominately various mammals. Birds and reptiles are used as preferred hosts by several Hyalomma species, and only on certain stage: adults of H. aegyptium parasitize tortoises; immature stages of H. marginatum parasitize birds. It is hypothesized that relationships of H. aegyptium adults (subgenus Hyalomma s. str.) with reptiles are secondarily in origin. Immature stages of H. aegyptium retain the primary wide diapason of hosts, which are various small mammals, birds and reptiles. The life cycle of this species is the three-host type that is considered as a primary type in ixodid ticks. A typical scheme of relationships with their hosts in all well-examined Hyalommina species has following features: the adult stage parasitize large and medium sized mammals, immature stages parasitize small mammals, three-host life cycle. A variety of preferred hosts and types of life cycle is observed in the subgenus Euhyalomma. All species of this subgenus can be arranged into two groups. In the first group, the immature stages infest only small mammals and birds, and the adults parasitize large mammals; this type of host preferences is probably primary host-parasite relationships of Hyalomma. This group includes: H. albiparmatum, H. asiaticum, H. excavatum, H. franchinii, H. impeltatum, H. impressum, H. lusitanicum, H. marginatum, H. nitidum, H. schulzei, and H. truncatum. Hyalomma marginatum and H. schulzei are two-host species; H. excavatum is two- or three-host tick. All the remaining species (except H. albiparmatum, which life cycle is unknown) are three-host ticks. In the second group, the immature stages as well as the adult stage parasitize large mammals. This group includes: H. dromedarii, H. anatolicum, and H. scupense. These species are two- or one-host ticks.     

Infraspecific variation in Cerastium alpinum s. lat. (Caryophyllaceae) in Central Norway

Within Cerastium alpinum s. lat., several taxa have been circumscribed. Three infraspecific taxa are currently recognized in Nordic literature, but the opinions have varied about their taxonomic levels and relations to each other. Populations of these three taxa — alpinum, glabratum and lanatum — were investigated in Central Norway to elucidate taxonomic relationships on a regional scale. Morphometric analyses (PCA ordination) based on two data sets; one with 61 characters, and one without the 35 indumentum characters, gave different results. The former partly support the use of a higher rank for glabratum , but this analysis may put too much weight on indumentum characters. The reduced data set gave a more continuous transition between the three taxa, with no special status for glabratum. Crossing experiments between the taxa indicated full cross compatibility as far as relative seed set was concerned. Finally, vegetation analysis (DCA ordination) indicated that the three taxa have somewhat different ecological demands. This suggests that they, in spite of hybridization abilities, normally are maintained as separate entities due to habitat differences. A delimitation of three infraspecific taxa at the same level within C. alpinum s. lat. is thus supported by this study. Whether these taxa should be treated as subspecies or varieties strongly depends upon the definitions of these categories. Viewed in a wide geographical context, however, the arguments for subspecific rank are strengthened.     

Exotic ectoparasites of ostriches recently imported into the United States

Eleven species of ectoparasitic arthropods were collected and identified from ostriches (Struthio camelus) recently imported into the United States from Africa and Europe. Four of these species are reported from ostriches for the first time. The parasites included adult hippoboscid flies (Struthiobosca struthionis) and 10 species of adult ixodid ticks (Amblyomma gemma, A. lepidum, A. variegatum, Haemaphysalis punctata, Hyalomma albiparmatum, H. luscitanicum, H. marginatum rufipes, H. truncatum, Hyalomma sp., and Rhipicephalus turanicus. As a result of these findings, the U.S. Department of Agriculture prohibited further importation into the United States of ostriches and other flightless birds on an interim basis.     

A morphometric analysis of Prunus spinosa, P. domestica ssp. insititia, and their putative hybrids in Denmark

The taxonomy of Prunus subgenus Prunus is notoriously problematic and reliable discriminating characters for species and subspecies identification are missing. Principal Component Analysis (PCA) was performed on different morphological data sets and useful diagnostic characters for Prunus spinosa and P. domestica ssp. insititia were found. Many traditionally used morphological characters were found unreliable. The hypothesis of widespread hybridization between Prunus spinosa and P. domestica ssp. insititia in Denmark were examined using PCA and HYWIN. It was found that hybrids between the two taxa are rare in Denmark.     

Reduced male fertility is common but highly variable in form and severity in a natural house mouse hybrid zone

Barriers to gene flow between naturally hybridizing taxa reveal the initial stages of speciation. Reduced hybrid fertility is a common feature of reproductive barriers separating recently diverged species. In house mice (Mus musculus), hybrid male sterility has been studied extensively using experimental crosses between subspecies. Here, we present the first detailed picture of hybrid male fertility in the European M. m. domesticus-M. m. musculus hybrid zone. Complete sterility appears rare or absent in natural hybrids but a large proportion of males (~30%) have sperm count or relative testis weight below the range in pure subspecies, and likely suffer reduced fertility. Comparison of a suite of traits related to fertility among subfertile males indicates reduced hybrid fertility in the contact zone is highly variable among individuals and ancestry groups in the type, number, and severity of spermatogenesis defects present. Taken together, these results suggest multiple underlying genetic incompatibilities are segregating in the hybrid zone, which likely contribute to reproductive isolation between subspecies.     

Experimental transmission of Crimean-Congo hemorrhagic fever virus: role of 3 vector species in the maintenance and transmission cycles in Senegal

In this article, we published the role of three species of ticks Amblyomma variegatum (Fabricius, 1974), Hyalomma marginatum rufipes (Koch, 1844) and Hyalomma truncatum (Koch, 1844) in the maintenance and transmission of the CCHF virus. The imagos of these species were infected by intracoelomic route. Vertical transmission (transtasial and transovarial) and horizontal transmission for different stases were studied by isolation on newborn mice, polymerase chain reaction, indirect immunofluorescence and ELISA. Our results proved that 15 days after inoculation, infection rates of 100% were noted with Hyalomma marginatum rufipes and Hyalomma truncatum. This rate is about 60% for Amblyomma variegatum. The imagos of the three species infected have transmitted the virus to their host during blood feeding (100%). A high transovarial transmission for Hyalomma marginatum rufipes and Hyalomma truncatum were observed (respectively 53 and 50%). This rate is about 12% for Amblyomma variegatum. The tick infection does not persist up to the first generation for the three species studied. Ticks are temporary reservoirs vectors but not permanent reservoirs of CCHF virus.     

The use of RAPD markers in Hordeum phylogeny.

The phylogenetic relationships among 39 wild Hordeum species, subspecies, and cultivated barley were investigated using RAPD markers as discriminating characters. Seventy-six RAPD fragments were generated using 12 single decameric primers of arbitrary nucleotide sequences. Amplification reactions resulted in fragments ranging in length between 200 and 2000 bp. Clearly resolved bands were scored for their presence or absence in a binary matrix. Amplified products were treated as independent characters to generate a phenogram using the NTSYS-PC package. Tree topology was generally found to be consistent with those based on morphological treatments. However, a few species like H. erectifolium, H. jubatum and, to a lesser extent, H. bulbosum occupied a position different from previous classifications. The results demonstrated that RAPD technology represents a useful and reliable tool for detecting polymorphism for phylogenetic studies. Key words : RAPD analysis, molecular markers, phylogenetic studies, Hordeum species, barley.     

The evolutionary history of sea barley (Hordeum marinum) revealed by comparative physical mapping of repetitive DNA

Background and Aims Hordeum marinum

is a species complex that includes the diploid subspecies marinum and both diploid and tetraploid forms of gussoneanum. Their relationships, the rank of the taxa and the origin of the polyploid forms remain points of debate. The present work reports a comparative karyotype analysis of six H. marinum accessions representing all taxa and cytotypes.

Methods

Karyotypes were determined by analysing the chromosomal distribution of several tandemly repeated sequences, including the Triticeae cloned probes pTa71, pTa794, pAs1 and pSc119·2 and the simple sequence repeats (SSRs) (AG)₁₀, (AAC)₅, (AAG)₅, (ACT)₅ and (ATC)₅.

Key Results

The identification of each chromosome pair in all subspecies and cytotypes is reported for the first time. Homologous relationships are also established. Wide karyotypic differences were detected within marinum accessions. Specific chromosomal markers characterized and differentiated the genomes of marinum and diploid gussoneanum. Two subgenomes were detected in the tetraploids. One of these had the same chromosome complement as diploid gussoneanum; the second subgenome, although similar to the chromosome complement of diploid H. marinum sensu lato, appeared to have no counterpart in the marinum accessions analysed here.

Conclusions

The tetraploid forms of gussoneanum appear to have come about through a cross between a diploid gussoneanum progenitor and a second, related—but unidentified—diploid ancestor. The results reveal the genome structure of the different H. marinum taxa and demonstrate the allopolyploid origin of the tetraploid forms of gussoneanum.     

Variations of the mandibular shape in extant hominoids: Generic, specific, and subspecific quantification using elliptical fourier analysis in lateral view

While a number of studies have documented the mandibular variations in hominoids, few focused on evaluating the variation of the whole outline of this structure. Using an efficient morphometrical approach, i.e. elliptical Fourier analysis, mandibular outlines in lateral view from 578 adult hominoids representing the genera Hylobates, Pongo, Gorilla, Pan, and Homo were quantified and compared. This study confirms that elliptical Fourier analysis provides an accurate characterization of the shape of the mandibular profile. Differences in mandibular shape between hominoid genera, species, subspecies, and to a lesser extent between sexes were demonstrated. Mandibles in great apes and hylobatids subspecies were generally less distinct from each other than were species. However, the magnitudes of differences among subspecies of Gorilla and Pongo approached or exceeded those between Pan troglodytes and P. paniscus. The powerful discrimination between taxa from the genus down to subspecific level associated to the relatively low level of intrageneric mandibular polymorphism in great apes provides strong evidences in support of the taxonomic utility of the shape of the mandibular profile in hominoids. In addition, morphological affinities between Pongo and Pan and the clear distinction between Homo and Pan suggest that the mandibular outline is a poor estimate of phylogenetic relationships in great apes and humans. The sexual dimorphism in mandibular shape exhibits two patterns of expression: a high degree of dimorphism in Gorilla, Pongo, and H. s. syndactylus and a relatively low one in modern humans and Pan. Besides, degree of mandibular shape dimorphism can vary considerably among closely related subspecies as observed in gorillas, arguing against the use of mandibular shape dimorphism patterns as characters in phylogenetic analyses. However, the quantification of the mandibular shape and of the variations among hominoids provides an interesting comparative framework that is likely to supply further arguments for a better understanding of the patterns of differentiation between living hominoids.     

Systematics of Pseudorchis albida s.l. (Orchidaceae) in Europe and North America

In Scandinavia Pseudorchis albida (Orchidaceae)usually divided into the lowland and subalpine P. albida s.s. and the more or less alpine P. straminea. There have been some uncertainties and conflicting views concerning die taxonomic treatment of diese taxa. To address this issue, herbarium specimens of P. albida s.l. were studied for variation in morphological characters. A small-scale population study approach was used, as herbarium sheets with two or three plants were used as population samples. Canonical Variates Analysis (CVA) indicated a distinction between taxa in population means, corresponding to P. albida s.s. and P. straminea , respectively. Principal Components Analysis (PCA), however, revealed an overlap between individuals of the two taxa. The PGA analysis, furthermore, revealed that the overlap was considerably larger in material from Central Europe man in material from Fennoscandia. Student t -tests on separate characters confirmed the picture, wim more characters significantly different in Fennoscandian than in Central European material. Furthermore, a Tukey-Kramer test revealed that there were small differences between regional populations of P. albida s.s. , while there were several significant differences in single characters between the Norm American regional population of P. straminea , as compared with the Central European and Fennoscandian regional populations. In Central Europe there is no clear separation between taxa, while in Fennoscandia the taxa are more clearly separated. This probably means that there is a difference in the time of establishment in the different regions. The author suggests a distinction of taxa at the subspecies level, and argues that the clear distinction seen in Fennoscandian material is due to separate immigration histories for die two subspecies into Fennoscandia after the last period of glaciation.     

A phylogenetic reassessment of Hyalenna Forbes and Dircenna Doubleday, with a revision of Hyalenna (Lepidoptera: Nymphalidae: Ithomiinae)

Abstract.  A morphological study of adults and immature stages and a cladistic analysis were conducted to clarify the relationships amongst members of the sister nymphalid butterfly genera Dircenna Doubleday and Hyalenna Forbes. Two species formerly included in Dircenna , Ithomia paradoxa Staudinger and Dircenna hugia Schaus, clustered with Ithomia perasippa , the type species of Hyalenna . Therefore, the first two species are transferred from Dircenna to Hyalenna ( comb.n .). Hyalenna and Dircenna as now conceived are both monophyletic and well supported by morphological apomorphies and branch support measures. The taxonomy and classification of Hyalenna are reviewed and seven species and twenty-four taxa are recognized, including one new species, Hyalenna buckleyi sp.n. , and eleven new subspecies: Hyalenna paradoxa incachaca ssp.n. , H. perasippa valencia ssp.n. , H. perasippa ortygiosa ssp.n. , H. perasippa solitaria ssp.n. , H. buckleyi pomacocha ssp.n. , H. alidella exsulans ssp.n. , H. alidella cinereola ssp.n. , H. alidella vesca ssp.n. , H. sulmona balsamica ssp.n. , H. sulmona hyalina ssp.n. and H. sulmona tersa ssp.n. A key for the identification of all taxa, and illustrations of male and female genitalia for all species and adult specimens of all taxa (where known), are presented. The natural history of each species is summarized and the immature stages of H. paradoxa and H. sulmona are described for the first time.     

早期形态发生:一种解决贻贝科(软体动物门:双壳纲)分类、系统发育和进化问题的方法

作者对贻贝科贝类的幼虫和幼贝期发育阶段形态结构的出现和变化顺序进行了研究,其约60个不同分类单元的个体发生可归纳为4种形态发生类型或模式。主要对3个形态发生区域的阶段形态结构的起源、发育变化和同源性做了研究。其一,即中央区域,开始形成于前双壳Ⅰ期(PD-Ⅰ),在某个分类单元它可以在前双壳Ⅱ期(PD-Ⅱ)和幼贝期(N)形成,而在其它分类单元则在前双壳Ⅱ期、幼贝期和双壳期(D)形成;第二区域,即背部后区,在幼贝期出现;第三区域,即背部前区,出现于双壳期。双壳期背部后区在某个分类单元起源于幼贝期的形态构造,在其它分类单元则可能起源于双壳期的形态构造。与在贻贝分类学上应用的成体特征相比,早期发育阶段中央和背部后区的形态结构显示出很明显的发育顺序或特征变化规律。根据以前人们熟知而尚未应用到分类和系统发育研究中的早期发育阶段形态特征,作者重新修订了Soot-Ryen的现生贻贝科种上阶元分类系统,重新提出了科内系统发育关系。修订的分类系统表明,Scarlato and Starobogatov(1984)提出的贻贝科各亚科由偏顶蛤亚科开始,沿4条系统发育路线演化发展,对应其早期发育阶段的4类形态发生类型或模式。     

Unraveling evolutionary lineages in the limbless fossorial skink genus Acontias (Sauria: Scincidae): are subspecies equivalent systematic units?

Subspecies in the limbless, endemic African fossorial skink genus Acontias constitute ill-defined operational taxonomic units, consequently considerable systematic debate has lingered on the systematic diversity within Acontias. In the present study, the systematic affinities among acontine taxa are explored with the utility of partial sequence data from two mitochondrial gene loci (16S rRNA and cytochrome oxidase subunit 1 (COI)) for all taxa, while two additional loci (12S rRNA, cytochrome b) were used to investigate relationships within the Acontias meleagris complex. Phylogenetic results, derived from the combined analysis, revealed two monophyletic clades. Clade 1 is comprised of small-bodied skinks while clade 2 comprised the medium bodied skinks. Within clade 2 none of the traditionally recognized subspecies formed reciprocally monophyletic groups. Furthermore, constraining the topology and enforcing sister taxa relationships between the assumed subspecies, consistently recovered a topology that was statistically significant worse, indicating that the traditionally designated subspecies groupings probably represent invalid taxonomic units, thus clearly reflecting considerable discord with current taxonomy. The burrowing life style of these lizards has probably led to marked convergent evolution and constrained the development of diagnostic morphological characters among these species. Morphological similarities in color as well as scale architecture within Acontias are labile and highly homoplaseous and do not reflect the evolutionary history of the group. Taxonomic implications of these results are discussed.     

Phylogenetics and the conservation of rare taxa in the Eucalyptus angustissima complex in Western Australia

Effective conservation that preservesevolutionary entities and processes requiresconservation action that is targeted atappropriate taxonomic units. Molecularphylogenetics can contribute to the resolutionof taxonomic uncertainties by defining therelationships between populations and species,and allowing the recognition of taxonomicentities that reflect evolutionary history.Some uncertainty surrounds the status of taxain the Eucalyptus angustissima complex,which includes threatened taxa with restricteddistributions. The phylogenetic relationshipsbetween populations and taxa in the E.angustissima complex were investigated usingnuclear and chloroplast RFLP analysis.Phylogenetic relationships confirmed thegenetic differentiation of two taxa, identifiedspecies level divergence between two taxacurrently recognised at subspecies level, andestablished the identity of a previouslyundetermined population. Phylogeneticrelationships between the taxa were notcongruent with presumed relationships based onmorphological characters alone.     

Asymmetrical reproductive character displacement in the house mouse

Our study addressed reproductive character displacement between two subspecies of the house mouse, Mus musculus musculus and Mus musculus domesticus that hybridize in Europe along a zone where selection against hybridization is known to occur. Based on a multi-population approach, we investigated spatial patterns of divergence of mate preference in the two taxa. Mate preference was significantly higher in the contact zone than in allopatry in both subspecies, suggesting that reproductive character displacement occurs. Moreover, patterns of preference were stronger in M. m. musculus than in M. m. domesticus, indicating an asymmetrical divergence between the two. In the context of selection against hybridization, our results may provide empirical support for the hypothesis of reinforcement in a parapatric hybrid zone. We discuss factors that could explain the asymmetrical pattern of divergence and the possible impact of a unimodal structure on the maintenance of premating divergence between the two subspecies.