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Co-occurring biodiversity and global heating crises are systemic threats to life on Earth as we know it, especially in relatively rare freshwater ecosystems, such as in Iran. Future changes in the spatial distribution and richness of 131 riverine fish species were investigated at 1481 sites in Iran under optimistic and pessimistic climate heating scenarios for the 2050s and 2080s. We used maximum entropy modeling to predict species' potential distributions by hydrologic unit (HU) occupancy under current and future climate conditions through the use of nine environmental predictor variables. The most important variable determining fish occupancy was HU location, followed by elevation, climate variables, and slope. Thirty-seven species were predicted to decrease their potential habitat occupancy in all future scenarios. The southern Caspian HU faces the highest future species reductions followed by the western Zagros and northwestern Iran. These results can be used by managers to plan conservational strategies to ease the dispersal of species, especially those that are at the greatest risk of extinction or invasion and that are in rivers fragmented by dams. [ABSTRACT FROM AUTHOR]

Iran is one of the world's fish biodiversity hotspots. Most riverine fish species in this country are currently under threat by human activities. In addition to those threats, climate change is expected to alter rainfall and temperature regimes, imposing further limitations, particularly to endemic fishes. Therefore, understanding how these species respond to climate change is an important issue in the conservation of Iranian freshwater biodiversity.This study aimed to predict the potential distribution of 16 endemic fish species under two climate change scenarios (i.e., RCP 4.5 and RCP 8.5) in river habitats of Iran in the 2050s and the 2080s. The analysis is conducted using the MaxEnt model. Seven environmental variables were used for modelling: maximum width, elevation, river slope, the basins occupied by the target species, average annual precipitation, annual mean temperature, and the temperature difference between the coldest and hottest months of the year.The validation of the model for each species showed that AUC (area under the ROC curve) scores range from 0.87 to 0.99. Under optimistic (RCP 4.5) and pessimistic (RCP 8.5) scenarios in the 2050s and the 2080s, species will face four different outcomes to their habitat range: reduction (three species), expansion (five species), reduction and expansion (seven species) and no change (one species).It is concluded that most species that are unique in their geographical area will face serious challenges from climate change. Delineating species range changes under climate change scenarios can help prioritise conservation measures for these valuable species. Considering our results, the translocation of species to new locations also should be investigated. [ABSTRACT FROM AUTHOR]

The Hyrcanian Forest holds broad leaf forest remnants dating back to the early Cenozoic Era, which once covered a vast area of the North Temperate Zone. Today, many rivers within this region have been altered by human activities and urgently need rehabilitation. In this regard, 35 wadeable rivers including 14 reference and impacted sites were investigated to determine how different human pressures altered riverine landscapes and habitats. Hence, five common human pressures (agriculture, urbanization, aquaculture, dams, aggregate mining) were identified, then the riverine landscape and habitat condition of each site were assessed. At each site, 17 aquatic, riparian, and terrestrial features, including abiotic and biotic substrate types, were investigated. The number and ratio of pressure-influenced channel features and substrate types differed from those in reference sites. Reference sites were dominated by microlithal, mesolithal, and macrolithal abiotic substrates and large wood, algae, and coarse particulate organic matter biotic substrates. Urbanized sites were most altered and dominated by single channels, steep unvegetated riprap banks, and algae substrate. The results provide valuable information for managers and decision-makers to restore riverine ecosystems considering the impaired parameters resulting from human pressures. [ABSTRACT FROM AUTHOR]

The active geology and ecological diversity in the Persian Gulf drainages create a rich source of biodiversity for the tooth‐carp fishes from which evolutionary processes can be studied. Here, we explored morphological differentiation in tooth‐carp Aphaniops hormuzensis from three ecologically diverse habitats (Khurgu sulphuric spring, Shur high‐salinity river and the urban canal) in the Hormuz riverine system, southern Iran. The evidence of sexual dimorphism was found in fish and otolith traits mainly in the Khurgu spring. The extreme conditions of the Khurgu spring could lead to the microhabitat segregation by fish individuals with different ages and spatial sexual segregation, and caused high intrapopulation differentiation in this site. However, the most diverged population belonged to the urban canal, which had more elongated pre‐ and post‐dorsal lengths, as well as a deep otolith. In sum, the observed phenotypic variations in A. hormuzensis are mainly influenced by sex‐dependent factors such as sexual dimorphism and the specific ecological condition of the Khurgu sulphuric spring (extreme sulphur‐rich habitat) and the urban canal (high food availability and hybridization event). This study provides the basic opportunity to explore the relative roles of ecological divergence and habitat differences in the morphological differentiation of A. hormuzensis. [ABSTRACT FROM AUTHOR]

The relationships between ontogenetic and sexual factors with otolith morphology are poorly studied in killifish genus Aphanius Nardo, 1827. We studied the ontogeny, morphological sexual and side dimorphism in the saccular otolith of a scaleless killifish Aphanius furcatus inhabiting a high‐salinity environment in Southern Iran. The results highlighted growth‐dependent variability in the otolith of A. furcatus, which might be useful to define its certain developmental stages. We underlined those characters that were consistent during fish development, which could be valuable to identify the species. Considering the ontogenetic pattern of otolith in A. furcatus, it was found that the otoliths of young individuals have largely been influenced by sexes and that it was reduced during the fish development. Morphological side dimorphism in the otolith of A. furcatus tends to be increased during fish development. The outcomes of this study are essential since they have provided new information on ontogenetic changes and the sexual and side dimorphism of Aphanius otolith. [ABSTRACT FROM AUTHOR]

Aphaniops dispar, widespread around the Arabian Peninsula, was recently separated in four species (A. dispar, A. hormuzensis, A. kruppi, A. stoliczkanus) by molecular results and colour patterns, but the morphological differences are small and call for more studies. Here we report differences in skeleton and median fin osteology of these species. In addition, we introduce the term 'modified caudal vertebra' to describe caudal vertebrae that are not directly associated with caudal ray support but are visibly modified from a 'usual' caudal vertebra. Aphaniops hormuzensis, an endemic species to southern Iran, has a significantly higher number of modified caudal vertebrae compared to the more widespread A. stoliczkanus and A. dispar, and also to A. kruppi. This is a surprising result as the caudal skeleton and related structures of the posterior caudal vertebral column have yielded successful results in separating between families or genera, but there are only a few studies that have examined these structures for their role in species diagnosis. Our study also highlights that state‐of‐the‐art methods in X‐raying and improved staining procedures assist in the discrimination of superficially similar species. [ABSTRACT FROM AUTHOR]

Despite their importance, our knowledge of the biodiversity and phylogeography of sponges in the Persian Gulf and adjacent waters is largely unexplored. Such knowledge, however, is necessary to better understand those patterns and processes that generated and maintained sponge diversity in this largely separated marginal sea, and to create a foundation for future exploration of the marine natural resources in this area. Recently, several expeditions resulted in the so far largest sponge collection of the Persian Gulf and Gulf of Oman, which now led towards the most comprehensive molecular taxonomic inventory of sponges in this region to date. We analyzed a total of 143 samples molecularly and identified 52 operational taxonomic units (OTUs) based on 28S and CO1. Additional molecular comparison with the Red Sea sponge fauna indicate a strong differentiation and high levels endemism between the Persian Gulf and the Red Sea respectively, supporting findings for other marine phyla. [ABSTRACT FROM AUTHOR]

The embryonic and early developmental stages of Killifish Aphanius hormuzensis reared in a laboratory condition were investigated in detail using light microscopy. The adult male and female of A. hormuzensis were captured from the wild. They were kept in laboratory condition for 2 months and spontaneously spawned at 28 ± 1°C. The fertilized macrolecithal eggs were characterized with spherical shape, transparent and average 1.60 ± 0.20 mm in diameter. The embryonic development was started with fertilized egg activation and the first cleavage was observed in 2.5 hr ± 20 min post fertilization (hpf), followed by blastula formation and gastrulation stages in 7 and 11 hpf respectively. First somite was observed in 20 hpf and the heart beating was detected in 25 hpf. The early ontogeny of A. hormuzensis was followed by complements of organs formation and the newly hatched larva with 4.9 ± 0.10 mm in length was detected in 166 hpf. This study provides a basis for further research on reproductive biology, conservation and breeding of this species. [ABSTRACT FROM AUTHOR]

Numerous riverine ecosystems in Iran are currently suffering due to various human activities, which are directly affecting the physico-chemical conditions of running waters and influencing the aquatic biota. In this study, we chose the cold-water streams of the Southern Caspian Sea Basin in Iran to analyse human pressures and develop a multi-metric fish index for the cold-water streams dominated by brown trout, Salmo trutta in this Basin. To do this, the fish and environmental data were sampled in 67 medium sized streams in the Caspian Mountains. In total, nine fish metrics were calculated. Moreover, 27 criteria describing major anthropogenic pressures for the entire sampling area were used to generate a regional pressure index that accounted for potential effects of multiple human pressures. For the index development, we first defined reference sites and then quantified differences of fish metrics between the reference and impacted sites. The multiple linear regressions were used for describing metric responses to natural environmental differences in the absence of any human pressures. Finally, by testing the ability of each metric for discrimination of impacted and reference sites as well as their sensitivity to a gradient of regional pressure index, the metrics density of brown trout and biomass of sub-adult brown trout explained human pressures best. Our multi-metric fish index performed well in discriminating between reference and impacted sites, giving a significant negative linear response to a gradient of regional pressure index. In general, the development of such an index offers an opportunity to enhance national bio-monitoring programmes in Iran. [ABSTRACT FROM AUTHOR]

Studies of the taxonomy and present-day distribution of landlocked populations of the killifish Aphanius Nardo, 1827 (Cyprinodontidae) provide a key to understanding their zoogeographic history, and shape conservation strategies and habitat management. Here we report for the first time on the sympatric occurrence of the rare Mesopotamian tooth-carp A. mesopotamicus with a member of the widely distributed common tooth-carp ( A. dispar group). Both were found in the Shadegan Wetland of the Mesopotamian drainage system (Southwestern Iran). External characters of individuals, otolith morphology and molecular data based on the cytochrome b gene confirm species identification. The otolith morphology of A. mesopotamicus , hitherto unknown, is strikingly similar to previously described otoliths of A . cf. pluristriatus from the Khonj stream (Southern Iran), which apparently reflects a close relationship between these recently diverged species. Several of the specimens recovered from the Shadegan Wetland showed intermediate characters in pigmentation, otolith morphology and some morphometric values. These specimens are interpreted as putative hybrids, which is additionally supported by a multivariate analysis of the morphometric data. Previous reports suggest that natural hybridization between species of Aphanius results largely from range overlap and range extension. A virtual distribution map derived from climatic modelling studies based on DIVA-GIS (7.5.0) indicates that sympatry of the A. dispar group and A. mesopotamicus is primarily determined by levels of precipitation during the warmest quarter, and is likely to occur only in Southwestern Iran. [ABSTRACT FROM AUTHOR]

Abstract: A primarily vicariance-based speciation has been suggested for the killifish genus AphaniusNardo, 1827, but ecological factors are likely to have promoted the speciation processes in addition. Here, we report on the discovery of a unique Aphanius species from Southern Iran and show that also regressive evolution has shaped the present-day diversity of Aphanius. The species is characterized by complete absence of scales and reduction in the biomineralization of hard structures, particularly of the caudal skeleton and jaw teeth. Based on mt-DNA sequences, morphometric and meristic data, osteology, jaw teeth and otoliths, it is described as Aphanius furcatus sp. n. The new species is sympatric with A. dispar (Rüppell, 1829) in salty rivers and hot sulphuric springs in the Hormuzgan Basin (Southern Iran), and is sister taxon to this species plus A. ginaonisHolly, 1929. Based on geological data we estimate that the divergence between the lineages of A. furcatus and A. dispar is about 12–14million years old. We conclude that the reductive phenomena observed in A. furcatus have evolved as an evolutionary response to the extreme habitat conditions in order to save energy (because storage of Ca2+ is not necessary), and to transport oxygen efficiently. The results confirm that regressive evolution is an important factor in speciation and occurs independently in separate lineages. [Copyright &y& Elsevier]

The Arabian cyprinodontid Aphanius dispar (Rüppell, 1829) is known to show considerable morphological variation. It has remained unknown, however, whether this variation is a result of environmental differences or allopatric divergence owing to geographical isolation. In this study, 11 populations of A. dispar from three geographically separated basins were analysed, that is, the Makran Basin (I, one river system), the Hormuzgan Basin (II, five rivers and three hot springs) and the Helleh Basin (III, two hot springs) in southern Iran. Statistical analyses do not indicate significant differences between the fishes from river and hot spring habitats ( T-test, p < 0.05), which is also supported by the Canonical Discriminant Analysis (CDA). Nevertheless, morphometric and meristic characters of the fishes, as well as otolith morphology and morphometry, demonstrate that six phenotypic characters discriminate the A. dispar populations of the three basins, that is, (1) predorsal distance (Prdd.SL), (2) head length (HL.SL), (3) pelvic fin length (Lplf.SL), (4) number of pelvic fin rays, as well as relative length of both the (5) medial part and (6) rostrum of the otolith. In addition, these characters display a consistent pattern of variation, thus providing support for the assumption that the phenotypically different A. dispar populations are a result of geographical isolation and not related to environmental differences. It is likely that the geological history of the drainage systems caused isolation event(s) that may date back to the Pleistocene (1.8 million years before present). The high phenotypic differences might suggest that the A. dispar populations from the three studied basins represent separate subspecies or even species. [ABSTRACT FROM AUTHOR]

A new species of tooth-carp, Aphanius arakensis sp. n., is described from the Namak Lake basin in Iran. The new species is distinguished by the congeners distributed in Iran by the following combination of characters: 10-12 anal fin rays, 28-32 lateral line scales, 10-13 caudal peduncle scales, 8-10 gill rakers, 12-19, commonly 15-16, clearly defined flank bars in males, a more prominent pigmentation along the flank added by relatively big blotches in the middle and posterior flank segments in females, a short but high antirostrum of the otolith that has a wide excisura, and a ventral rim with some small, drop-like processes, and 19 molecular apomorphies (17 transitions, two transversions) in the cytochrome b gene. It was suggested based on the phylogenetic analysis that the new species is sister to A. sophiae from the Kor River and that A. farsicus from the Maharlu Lake basin is sister to A. arakensis plus A. sophiae. A noticeable feature of the Aphanius diversity in Iran is the conservatism of the external morphology as well as morphometric and meristic characters, while distinctive differences are present in genetic characters, otolith morphology, and male color pattern. Transformation of the latter was probably driven by sexual selection. [ABSTRACT FROM AUTHOR]

Aphanius Nardo, 1827 (Actinopterygii, Cyprinodontidae) is a widely distributed genus in the Mediterranean and Persian Gulf area and includes several endangered species. The otolith morphology in Aphanius is known to represent a valuable tool for the taxonomy, and is also indicative for the genetic diversity of a particular population. The present study focuses on the otoliths of the endangered A. ginaonis (Holly, 1929), which is endemic to the Geno hot spring in southern Iran. The taxonomic status of A. ginaonis has repeatedly been questioned, and some scholars have argued that it merely represents a morphological variation of the widespread A. dispar. We present a comparison of the otolith morphology of A. ginaonis (52 specimens) with that of A. dispar (Rüppell, 1828) from the Mehran River Basin (southern Iran) (17 specimens) and an A. dispar population from the Persian Gulf coast of the United Arab Emirates (32 specimens). Our data obtained from SEM pictures, otolith morphometry and statistical analyses suggest that A. ginaonis represents a valid species. In A. ginaonis individuals with a standard length exceeding 23 mm, the otolith variables length–height and rostrum length represent useful complementary diagnostic characters discriminating this species from other Aphanius species. Besides ontogenetic variation, we found extremely high otolith form variability in A. ginaonis, including some otoliths with a morphology distinctly deviating from the basic morphology type. We hypothesize that these variations may be a result of the artificial introduction of A. dispar into the Geno hot spring during the last years and subsequent hybridisation. [ABSTRACT FROM AUTHOR]

Aphanius hormuzensis is an endemic tooth-carp found in the Hormuzgan drainage in S-Iran. This study aimed to investigate the size-dependent alternations of scale surface ornamentation in this species by conducting scanning electron microscopy and morphological analysis. A total of 50 wild fish individuals were captured from Shur River, and were classified into five size classes based on the standard length; SC-I (SL = 10-20 mm), SC-II (SL = 21-30 mm), SC-III (SL = 41-50 mm), SC-IV (SL = 51-60 mm), and SC-V (SL = 61-70 mm), and their scales were removed from below the dorsal fin (key scale) and caudal peduncle regions. The results revealed a clear trend of scale structural development in A. hormuzensis. The scale of small-sized (TL < 30 mm) and large-sized fishes (TL > 30 mm) differed, respectively, in the following characters; the overall shape (often circular vs. polygonal), relative focus size (large, FL/SL = 440-610 μm vs. intermediate and small, FL/SL = 100-330 μm), types of radii (only primary vs. three types), relative radii length (short, RL/SL = 100-180 μm vs. long, RL/SL = 320-450 μm), lepidont (absent or undeveloped vs. present and developed), and the relative lepidont length (short, LL/SL = 0.83-0.90 μm vs. intermediate and long, LL/SL = 1.2-2.2 μm). These character alternations could explain certain developmental stages in this species. The size-dependent changes in the surface micro-ornamentations as shown in this study suggest that these characters should be used cautiously for taxonomic studies of the aphaniid fishes., (© 2021 Wiley Periodicals LLC.)

To study scale based phylogenetic affinity, the ultrastructure and ornamentation characteristics of body key scales were studied for 12 gobiid species from the Iranian coast of the Persian Gulf including Qeshm and Hormuz Islands and the Makran coast of the Oman Sea using scanning electron microscopy (SEM) technique. The scales were removed from below the first dorsal fin, cleaned in potassium hydroxide solution 1%, and were prepared for the SEM imaging. The presence of both ctenoid and cycloid scales in the studied gobiids was revealed. The focus of ctenoid scales was positioned posteriorly, while the focus of cycloid scales was positioned in the postero-central part of the scale. In all the studied species, radii were located only on the anterior part of the scale, and the primary radii were dominant. Also, there were no granules in the inter circular space, but bifurcation was observed in some circuli. In most species, the teeth-like structures called lepidonts were present on the crest of circuli, which functionally help to firmly attach the scales to the epithelium. The dendrogram of the between-groups-linkage method sorted the gobiid species into the two main groups of five distinct clusters: (a) Cryptocentroides arabicus and Cryptocentrus cyanotaenia (the Cryptocentrus-lineage); (b) Bathygobius meggitti and Bathygobius cocosensis (the Glossogobius-lineage); (c) Coryogalops adamsoni and Coryogalops tessellatus (the Gobius-lineage); (d) Acentrogobius dayi, Istigobius ornatus, Favonigobius reichei, Aulopareia ocellata, and Silhouettea ghazalae (the Gobiopsis-lineage). It seems that the dendrogram topology obtained based on the macro-and microscopic structures of scales, reveals phylogenetic lineages of gobies that have already been proposed for these taxa. Hence, the results of this study are largely consistent with the previous molecular studies on the gobiid fishes and implied that besides other data, the analysis of scale shape and scale-surface microstructures could be served to study the diversification of gobiid species., (© 2020 Wiley Periodicals LLC.)

The fishes, which have currently named Aphanius Nardo, 1827 are the relict of the ancient ichthyofauna of the Tethys Sea. For a long time since 1827, the genus name has been subjected to revision by several researchers using mainly morphological features. Until recently, no comprehensive single- or multi-locus DNA barcoding study has been conducted on whole members of the family Aphaniidae. In the present study, by applying four conceptually different molecular species delimitation methods, including one distance-based method, one network-based and two topology-based methods, we examined a single-locus DNA barcode library (COI) diversity for the 268 sequences within the family Aphaniidae from the Old World (57 sequences are new in the present study and 211 sequences were retrieved from NCBI database). The molecular analyses revealed a clearer picture of intra-family relationships and allowed us to clarify the generic names, and also describe a new genus for the family Aphaniidae. Results supported distinction of three major clades related to three genera within this family: i) the first clade includes the A. mento group which are placed in a new genus, Paraphanius gen. nov., found in the Orontes (= Asi) and Tigris-Euphrates River drainage, the Levant in coastal waters and the Dead Sea basin, western Jordan, and in southern Turkey in the Mediterranean basins as well as in central Turkey. This clade positioned at the base of the phylogenetic tree, (ii) the second clade contains the A. dispar-like brackish water tooth-carps which are transferred to the genus Aphaniops Hoedeman, 1951 (type species, Lebias dispar), distributed in the coastal waters around the Red Sea and the Persian Gulf basins; and (iii) the third clade, the genus Aphanius Nardo, 1827 (type species Aphanius nanus = A. fasciatus) contains all the inland and inland-related tooth-carps, which are mainly distributed in the inland waters in Turkey and Iran and also in the inland-related drainages around the Mediterranean basin., Competing Interests: The authors have declared that no competing interests exist.

Paraschistura kermanensis from the endorheic Kerman-Naein basin in Iran is distinguished from its closest congeners (P. abdolii, P. delvarii, P. kessleri and P. naumanni) by lacking scales on the flanks anterior to the dorsal-fin origin, having scales on the caudal peduncle, and having the pelvic-fin origin situated anterior to or below the vertical through the dorsal-fin origin. It is also characterised by six fixed diagnostic nucleotide substitutions and a K2P nearest neighbour distance of 4% to P. abdolii in the mtDNA COI barcode region.

Two new species of Aphanius are described from the Kol drainage in southern Iran (Aphanius darabensis n. sp.) and the endorheic Kavir Basin in northern Iran (A. kavirensis n. sp.), and compared with eight closely related species. Aphanius darabensis n. sp. is sister to A. shirini, from which it is distinguished by molecular characters (cytochrome b) and the combination of three morphological characters: 9-18 flank bars in males (vs. 7-10), females with irregular vertical patches of brown color on the flank (vs. prominent dark brown blotches of round or irregular shape), and symmetrically-shaped triangular to trapezoid otoliths with a rostrum distinctly longer than the antirostrum (vs. quadrangular to trapezoid otoliths with short and equally sized rostrum and antirostrum). Aphanius kavirensis n. sp. is closely related to a group containing A. sophiae, A. mesopotamicus and A. pluristriatus, from which it is distinguished by cytochrome b characters and the combination of three morphological characters: females with irregularly arranged large blotches of dark brown color on the flank, short pectoral fin in both sexes (13.4-18.1% SL in males, 11.2-18.3% SL in females), and asymmetrically shaped triangular to trapezoid otoliths with a pronounced predorsal region. Our tree based on the cytochrome b data demonstrates that the Iranian inland and inland-related Aphanius species (IIRAS) form a monophyletic clade with three subclades (A. vladykovi -, A. shirini -, A. sophiae subclades). The A. sophiae subclade, which is the most diverse of the three subclades, can be further divided into three lineages (A. isfahanensis -, A. farsicus -, A. sophiae lineages). The temporal diversification of the IIRAS clade is discussed and two evolutionary groups of Aphanius are depicted. Aphanius vladykovi, together with A. shirini, A. darabensis n. sp. and A. isfahanensis characterize the "old" evolutionary group. Their divergences may have happened 10-5 m.y. ago (Late Miocene-Early Pliocene). Aphanius farsicus, A. arakensis, A. sophiae, A. mesopotamicus, and A. pluristriatus, together with Aphanius kavirensis n. sp., represent the "young" evolutionary group, which developed in the Late Pleistocene (100,000-11,700 y. ago) and Early to Middle Holocene (c. 11,700-4,000 y. ago).

A substantial number of species within the tooth-carp Aphanius Nardo, 1827 (Cyprinodontidae, Teleostei) has been recorded from the endorheic drainage systems of Iran, and several isolated populations in these systems may deserve species status. Descriptions of these species and populations have been based mainly on morphological and molecular data; however, the characters related to the fish scales have not up to now been intensively studied and employed for the identification of the species belonging to this genus. The objective of this study is to test as to whether (i) scale surface morphology, (ii) scale surface microstructure, and (iii) scale size can be used to discriminate species and/or populations and, (iv) to discuss the possible causes of the observed differences. To achieve these objectives, scales of three species of the genus Aphanius from endorheic basins in SW Iran, viz. A. sophiae (nine populations from the Kor River Basin), A. farsicus (four populations from the Maharlu Lake Basin) and A. pluristriatus (a single population from the Mond River Basin) have been studied using SEM images, scale measurements, and uni- and multivariate statistics. It is opined that scale surface morphology and microstructure cannot help in distinguishing the species, but can be employed to discriminate certain populations of A. sophiae (those from Safashahr, Kharameh, Tashk, Gol). In addition, scale size and J-indices, respectively, represent a valuable tool for species separation, which corroborates earlier studies for the use of these indices in taxonomy. Major driving forces of the differentiation within A. sophiae probably include habitat fragmentation resulting from the geological history and local adaptations. Thus the differentiation results from a balance between both genetic and environmental effects.