Western Transcaucasia Research Articles

Historical drivers and future streamflow variations of the Kura River in the Western Transcaucasia region of Georgia: Analysis of tree-ring chronologies from 1720 to 2021 CE

In this paper, we conduct a dendrochronological study in the Western Transcaucasia region of Georgia to elucidate the relationship between tree growth and climatic variables (temperature, precipitation, and streamflow) in order to reconstruct the paleohydrology of the Kura River and to assess the impacts of climate change on water resources. We analyzed tree-ring chronologies from 1720 to 2021 CE, and analysis of July–August streamflow revealed five wet phases (1739–1753, 1764–1807, 1811–1831, 1962–1988 and 2000–2019 CE ranging in duration from 11 to 49 years) when streamflow exceeded the long-term mean, and four dry phases (1720–1738, 1832–1881, 1936–1961 and 1989–1999 CE) when streamflow was below average. The most extreme wet and dry years were 1771 and 1851 CE, respectively. Spatial correlation patterns of Kura River reconstructed runoff against sea level pressure (SLP) are negative in the Arctic and positive in mid- to high-latitude Eurasia. The Arctic Oscillation (AO)/North Atlantic Oscillation (NAO), and solar activity are important drivers of hydroclimatic conditions in the Mediterranean region, by modifying the location of high-pressure areas and large-scale atmospheric circulations. Comparison of our Kura streamflow reconstructions and data in the Old World Drought Atlas (OWDA) reveals a moderately positive correlation (0.40) throughout the study period, being strongest over the Southern Caucasus and Eastern Turkey. This validates the drought signals and emphasizes the importance of using multi-proxy approaches, supported by spatially resolved data, to enhance understanding of regional hydroclimatic variability and drought patterns. We additionally conducted future streamflow scenarios, most notably under the high-emission SSP585 scenario, in line with global trends in CMIP6 projections. Our streamflow results indicate a trend towards increasing drought severity, which highlights the need for urgent adaptation and mitigation strategies in water resource management in this region. Ultimately, this study provides valuable information on historical hydroclimatic conditions in the Kura River Basin, which help to develop strategies to mitigate the hydrologic impacts of climate change in the Western Transcaucasia Caucasus region. We further highlight the importance of regional and global teleconnections for understanding regional hydrological dynamics.

The Caucasus and the Middle East in the early Holocene (According to recent archaeological research)

The article provides a comprehensive account of two significant prehistoric archaeological sites located in Adjara - Kobuleti village and Khutsubani, including their history, modern investigations, and findings. The authors highlight that bone remains were not uncovered during the excavation of Stone Age archaeological sites due to the soil'speculiarities. These remains could have shed light on the hunting environment of that era. Stone artifacts provide insight into the lives of ancient hunter-gatherers in our regionThus, a thorough examination of the stone industry is very important. According to this analysis the authors, a fascinating conclusion is drawn regarding the origins of the Western Transcaucasia stone industry. During the early Holocene era, a new method of stone processing and various types of tools emerged within the aforementioned region. This innovative approach was originally developed in the territory of Iran and Iraq. It's a Mlefaatien culture, which comprises of many significant sites. After conducting a techno-typological analysis of the stone complex, it is believed that hand pressure techniques, backed microblades, Kashkashok side-blow blade-flakes, grooved tools etc. emerged in Western Georgia result of the great migration process from Middle East in the beginning of the 10th millennia BC. This view is supported by the complete range of precise dates, which accurately reflect the migrations ways and times of the early migrants. Additional inquiries in this field, conducted in the future, will undoubtedly uncover even more fascinating insights.

Phylogeography of Oak Species in the Caucasus Based on the Results of Chloroplast DNA Analysis

In order to study the biogeography and history of the temperate biota of the Caucasian region, one of the centers of biological diversity of Eurasia, the phylogeographic structure of the chloroplast DNA of roburoid oaks (Quercus L. sect. Quercus, Fagaceae) was studied. We analyzed 926 trees of five species (Quercus robur, Q. petraea, Q. pubescens, Q. hartwissiana, Q. macranthera) from 70 populations located in different parts of the Caucasus. Sequencing of five fragments with a total length of more than 10 000 bp revealed eight haplotypes, which were typed using chloroplast microsatellite loci (cpSSR), sequencing, and restriction analysis. The Quercus phylogenetic tree, which includes 34 chloroplast haplotypes of Western Eurasian and East Asian roburoid species, confirmed the monophyly of roburoid oaks, which form several lines that do not have supported topological relationships with each other. The haplotypes identified in the Caucasus belong to two divergent Western Eurasian lineages. These haplotypes are endemic for the Eastern Black Sea and Caucasian region and do not have related haplotypes beyond its borders, which indicates the long-term presence of oaks in the Caucasus and the historical links of the oaks of the Caucasian region with the Eastern Crimea and Eastern Turkey in the absence of chloroplast DNA exchanges with the Eastern European part of the range. The results of the present study indicate that Caucasian populations could not significantly contribute to the colonization of northern territories, and were not influenced by more northern populations, at least through seed transfer. The distribution zones of the Caucasian haplotypes almost do not intersect with each other, occupying certain geographical areas, probably due to the settlement from individual glacial refugia and genetic drift. The geographical structure of chloroplast DNA variability indicates a long-term presence of oak in addition to Western Transcaucasia (Colchis refugium), in the regions of the Northwestern, Central, Eastern Caucasus and the Lesser Caucasus. The coincident composition of the haplotypes of the studied oak species within the geographic regions indicates the long-term coexistence of different species in different parts of the Caucasus.

Euomphalia schileykoi sp. nov. (Mollusca, Gastropoda, Helicidae) in the Transcaucasia

The study of the anatomy of Euomphalia aristata from the Western Transcaucasia showed that these populations differ from other species of the genus and from the typical populations of E. aristata from the North Caucasus in anatomy of the genitalia. E. aristata in Transcaucasia is characterized by the separation of the distal parts of the male and female sections of the reproductive system, a long oviduct, a very short proximal part of the vagina, and the location of the vaginal appendages close to the mucous glands. In E. aristata from the North Caucasus (North Ossetia), the male and female parts of the genitals are not separated in the distal part, the oviduct is short, the proximal part of the vagina has an S-shaped bend, and the vaginal appendages are located at a noticeable distance from the mucous glands. On these grounds we distinguishE. aristata in Transcaucasia as a separate species.

CONNECTIONS OF THE KUBAN LATE BRONZE AGE CENTER OF METALLURGY WITH THE CENTRAL AND WESTERN TRANSCAUCASIA

CONNECTIONS OF THE KUBAN LATE BRONZE AGE CENTER OF METALLURGY WITH THE CENTRAL AND WESTERN TRANSCAUCASIA

Russian Customs System and Its Functioning in Western Transcaucasia (early 19th century – 1861)

Russian Customs System and Its Functioning in Western Transcaucasia (early 19th century – 1861)

Revealing the stygobiont and crenobiont Mollusca biodiversity hotspot in the Caucasus: Part III. Revision of stygobiont microsnails (Mollusca: Gastropoda: Hydrobiidae) from the Russian part of Western Transcaucasia, with the description of new taxa.

A taxonomic revision of the stygobiont microsnails from north-western Transcaucasia (Krasnodar Krai, Russia) is presented. Two new generaSchapsugia gen. n. and Tachira gen. n. are established. It is shown that the genus Paladilhiopsis does not occur in this area, and the snails previously assigned to this genus belong to Schapsugia gen. n. Two nominal species Paladilhiopsis orientalis Starobogatov, 1962, and P. subovata Starobogatov, 1962 are considered junior synonyms of Sch. pulcherrima (Starobogatov, 1962), based on a re-examination of their conchological variation. In addition, two more species of Schapsugia are described: Sch. kudepsta sp. nov. and Sch. occultata sp. nov. The anatomical structure of another species, Geyeria valvataeformis Starobogatov, 1962 was studied for the first time. It is revealed that it should be placed in a separate genus (Tachira gen. n.). Thus, the previously proposed placement of the microsnails from north-western Transcaucasia in hydrobiid genera from the Balkan Peninsula (Geyeria, Paladilhiopsis) is rejected.

Մեծ Բրիտանիայի հակահայ քաղաքականությունը Ղարաբաղի հարցում (1918–1920 ԹԹ.) և արդի իրավիճակը

The Armenian people were hopeful that after the capitulation of Turkey in the First World War (WWI) and the dislocation of the Allies troops in Transcaucasia, territories of Western Armenia and Transcaucasia would be included within the First Republic of Armenia. However, the Armenian people experienced another disappointment. In 1918–1920, Great Britain, who had gained dominance in Transcaucasia, ignoring the promises given to the Armenians during the war and their particular contribution to the victory of the Allies, adopted anti-Armenian attitude towards the Karabakh conflict. By constant pressure on the Republic of Armenia and Artsakh, the British strived to achieve hegemony of Azerbaijan over Karabakh, nevertheless, the uncompromising stance and unwavering resistance of the Armenians of Artsakh thwarted the anglo-azerbaijani plan. The anti-Armenian attitude of Great Britain was shaped by its strategic plans. Its main purpose in Transcaucasia was to gain control over the economic and military-political life and withdraw Russia from the region and the East in general. With the aim to carry out this important plan, it chose as an ally its recent rival in e WWI – Turkey, and Azerbaijan, which was created with the latter’s help. This selection was influenced by their military-economic potential and anti-Russian attitude. Nowadays (since 1988) when the Armenians of Karabakh struggle for their reunion with Armenia, from which Karabakh was cut off and annexed to Azerbaijan with the help of Soviet Russia on July 5, 1921, the British adopted anti-Armenian attitude towards the Karabakh conflict once again. They supported the aggressive actions of Azerbaijan and its ally Turkey by all means. The United Kingdom’s attitude proceeds from its political and economic interests. The United Kingdom has a large share of Azerbaijan’s most profitable oil mines, as well as the Baku–Tbilisi–Ceyhan oil pipeline, which was put into operation in 2006 due to its active efforts (30% of the shares in the pipeline are owned by British Petroleum). The anti-Armenian and pro-Azerbaijani policy of Great Britain activated during the Second Karabakh War, which began on September 27, 2020, by Azerbaijan’s aggression against the Nagorno-Karabakh Republic (the Republic of Artsakh). British companies expressed their support to Azerbaijan during this war have signed new economic agreements as well as have provided the other side with the maps of mined areas. In October, 2020, during the session in the UN Security Council, the UK suspended resolution of the OSCE Minsk Group member countries with an appeal to stop the war. Hence, the UK supported both war, intervention of the third party countries and terroristic groups in it. Thus, England once again betrayed democracy, justice, right of nations to self-determination, though the United Kingdom publicly presents itself as a protector of these values.

Dispersal history of the golden jackal (Canis aureus moreoticus Geoffroy, 1835) in Europe and possible causes of its recent population explosion.

BackgroundData on the historical distribution of the golden jackal in Europe and its primary habitats are scarce. There are many new data on the population explosion and the rapid spread of the in Europe. However, the main factors for this expansion, the core population and its routes of dispersal, remain controversial or insufficiently studied.New informationThis study provides a profound analysis of the history of the jackal’s (Canis aureus moreoticus Geoffroy, 1835) occurrence in Europe, the factors limiting or those triggering its expansion on the continent. The analysis shows that the timing of the species appearance in Europe still remains unclear. Historical data show that the species is a typical inhabitant of South-Eastern Europe, with some pulsations within its core area, as well as extensions to the north and west of it in favourable periods. Nowadays, the increase of the species range in Europe is the largest documented population explosion on the continent. We argue that this expansion originates from only three core populations, the Peri-Strandja area and the Dalmatian coast in the Balkans and the east parts of Western Transcaucasia in the Caucasus. This population explosion is largely due to a unique combination of factors of an anthropogenic nature.

Mitochondrial phylogeny and biogeography of the genus Gobio (Teleostei: Cyprinidae) in Turkey

Species of the cyprinid genus Gobio Cuvier, 1816 are widely distributed in freshwater lakes and rivers in Turkey, which is a hotspot for freshwater fish diversity and endemism. The mitochondrial 16S, coxI and cytb genes were sequenced for 217 individuals representing 15 species of Gobio from Turkey. A total of 23 haplotypes were identified for each mitochondrial gene. The genetic distance matrices show that Turkish Gobio species are clustered into three groups (northwestern, central and northeastern Anatolia). Phylogenetic trees constructed with combined dataset by using maximum parsimony, maximum likelihood and Bayesian inference methods revealed that the Turkish Gobio species belongs to three well-supported groups in accordance with their geographic distribution: Group I comprises species found in Central Anatolia, Group II species ranging from eastern Thrace to western Anatolia (northwestern Group) and Group III contains only one species from Western Transcaucasia (northeastern Group). The estimated divergence times between the three Gobio groups, calculated using a conventional 1% rate of mutation for a fish mitochondrial cytb gene per million years, coincide with the late Miocene period in which the tectonic uplift of Anatolia and global climate fluctuations occurred. The relatively low genetic distance between Gobio species in the Turkish Lake District indicate that they are not good species.

Taxonomic review of the genus Calyptopsis of Turkey and adjacent areas (Coleoptera: Tenebrionidae: Tentyriini)

Abstract The taxonomic review of the tenebrionid genus Calyptopsis Solier, 1835 (Tenebrionidae: Tentyriini) of Turkey and Western Transcaucasia is given. Four new species and one new subspecies are described: C. capnisiformis volcanica subsp. nov. (from calderas of extinct volcanoes in Karaman and Manisa provinces), C. fouquei sp. nov. (Artvin Province), C. egecemi sp. nov. (Izmir Province), C. kaszabi sp. nov. (Bayburt Province), C. ottoi sp. nov. (Turkey: Hakkari and Van provinces; NE Iraq: Zagros Mts.). The lectotype of Calyptopsis lineimargo Reitter, 1897 is designated. The key and images for the Turkish and Western Transcaucasian species are given. In total, eleven species and one subspecies are known from Turkey, of which eight taxa are endemic for Anatolia.

The genus Parmelina (Parmeliaceae) in the Russian Caucasus

The present study provides a brief survey of the genus Parmelina in the Russian Caucasus (Western Transcaucasia, Western, Central and Eastern Caucasus) with a special emphasis on species distribution. The study area is located in the largest nature reserves of the Russian Caucasus — Kavkazskiy, Utrish, Severo-Osetinskiy, Erzi. This paper contributes to a better understanding of the genus in the Caucasus, focusing mainly on the author’s collections in different regions of the Northern Caucasus in 2008–2016. Four species have been identified from the Russian Caucasus — Parmelina carporrhizans, P. pastillifera, P. quercina and P. tiliacea. Notes on their morphology, anatomy, ecology and distribution are based on data obtained from a study of ca. 20 specimens of each species. The presence of whitish macules on the upper surface of P. carporrhizans (after Clerc, Truong, 2008) as an important diagnostic feature was shown. The Caucasian material shows the differences in size and proportions of ascospores, size and shape of pycnoconidia between critical species P. carporrhizans and P. quercina. An identification key for Caucasian Parmelina species is composed for the first time and can be used for central regions of Russia.

ECOLOGICAL AND ZOOGEOGRAPHICAL ANALYSIS OF CAPSID BUG FAMILY (INSECTA, HETEROPTERA, MIRIDAE) OF THE CAUCASUS

The aim is to generalize the scientific researches on the species composition of Hemiptera within the Caucasus and possible ways of its formation. Materials and methods. As research materials we used the collections of Hemiptera of the Zoological Institute of the Russian Academy of Sciences, the Institute of Zoology of the National Academy of Sciences of Azerbaijan as well as own collections and analysis of the available literature data. As the basic tool, we used the method of zoogeographical and ecological analysis of capsid bugs of the Caucasus. We also studied the hemipterofauna of the Mirid bugs of the individual regions of the Caucasus. Findings. The paper presents an analysis of the representation and relation of zoogeographical complexes of Hemiptera insects of Miridae (Heteroptera) family in the vast territories of the North Caucasus and Transcaucasia. The analysis confirmed the major conventional views on the possible ways of forming a complex of Mirid bugs in the Caucasus, although so far the issue has not been studied completely. Conclusion. Based on the analysis of faunal and taxonomic studies, we studied the main features of the zoogeographic structure of the most numerous family of Hemiptera in the region. The hemipterofauna of the Western Transcaucasia is dominated by the boreal complex, and the Transcaucasia is dominated by the Mediterranean complex with a significant part of the Iran-Turan. Currently, this family consists of around 600 species. Among Mirid bugs, cosmopolitans in this region are rare. In the North Caucasus, there are almost no endemic species, while in the South Caucasus, they account for about 5% of the total. The role of human factor in the formation of certain complexes is quite high.

Geographical features of the foreign electricity trade in the post-Soviet space

In recent years, many studies on the foreign electricity trade in the post-Soviet space have been published, but they have not considered the regional specific nature of this phenomenon. The aim of this work is to identify the regional features of development and degradation of foreign trade in the post-Soviet space over the period 1990–2015. The work is carried out using comparative-geographical, historical and evolutionary, and cartographic methods. The directions of construction and dismantling of the cross-border PTLs, cross-border power transmission, and the loading level of cross-border PTLs are determined. Analysis of these parameters has shown that the general trend for the post-Soviet space has become degradation of the foreign electricity trade and the service infrastructure despite the high potential for international cooperation in this industry. The exceptions are the Baltic states, which have established integral power systems due to unification of the electricity market parameters. The imbalances associated with the closure of the Ignalina nuclear power plant and active development of wind power plants, as well as the requirements of all-European policy, have facilitated construction of cross-border PTLs in the Baltic states. Assessment of prospects has shown that, provided the solution of some foreign policy issues, it is possible to create power pool systems based on the use of electricity produced by hydroelectric power plants in eastern Central Asia and western Transcaucasia. Intensification of electricity exports to Asian countries (China, Afghanistan, Iran, and Turkey) is typical of many countries. The main precondition for the development of foreign trade is an imbalance between growing power consumption in some regions and the availability of cheap energy in others. Less important are fluctuations in the currency rate, foreign policy phenomena, and the policy for developing national power systems.

First finds of Acalypha australis L. and Euphorbia maculata L. (Euphorbiaceae) in Dnipropetrovsk region

First finds of Acalypha australis L. and Euphorbia maculata L. (Euphorbiaceae) in Dnipropetrovsk region

Contribution to the lichen flora of the Shakhe River valley (Krasnodar Territory, Western Transcaucasia)

A total of 138 species of lichens, lichenicolous and allied fungi are reported from broad-leaved forests with Buxus colchica from the Shakhe River valley in the Caucasus State Nature Reserve (Krasnodar Territory, Western Transcaucasia). The list includes 10 foliicolous lichens, namely Arthonia microsticta, Bacidina apiahica, Byssoloma leucoblepharum, Fellhanera bouteillei, F. viridisorediata, Gyalectidium caucasicum, G. setiferum, Phyllogyalidea phyllophila, Porina oxneri, Strigula buxi. On the thallus of the latter, a lichenicolous fungus Lambinonia strigulae was found. Nine species are recorded for the first time for the Russian Caucasus, five of them being new for the Caucasus. Endococcus ramalinarius, Eopyrenula septemseptata and Pyrenula subelliptica are reported for the first time for Russia. The list includes five species included in the Red Data Book of Russian Fеderation (2008) and ten species included in the Red Data Book of Krasnodar Territory (2007).

History in the Context of the Georgian-Abkhazian Conflict

The 2014 disturbances in the Ukraine occasioned renewed discussion of the 2008 Russo- Georgian War. As the situation continued to worsen in eastern Ukraine, US President Obama announced on a visit to Poland at the start of June that the US and NATO would strengthen ties even with the non-NATO-member-states of the Ukraine, Moldova and Georgia. This last has aspirations of membership, even though it does not control the republics of South Ossetia and Abkhazia, which most of the world nevertheless regards as integral parts of Georgia. As long as the Georgian-Abkhazian dispute remains unresolved, there will be problems regarding inter-state relations with/for western Transcaucasia. And there can be no resolution of the Abkhazian issue without a proper understanding of Abkhazia’s history (both ancient and more recent); it was to try to ensure that the debate is not based on misconceptions, unsubstantiated assertions or even plain errors that this article was written. It is grounded on a consideration of a range of materials (from Agathias’ Greek text through relevant discussions in Georgian, Russian and English). The toppling of Abkhazia’s democratically elected president (Aleksandr Ankvab) at the end of May 2014 makes the question of Abkhazia even more topical.

Mitochondrial phylogeography, contact zones and taxonomy of grass snakes (Natrix natrix,N. megalocephala)

Grass snakes (Natrix natrix) represent one of the most widely distributed snake species of the Palaearctic region, ranging from the North African Maghreb region and the Iberian Peninsula through most of Europe and western Asia eastward to the region of Lake Baikal in Central Asia. Within N. natrix, up to 14 distinct subspecies are regarded as valid. In addition, some authors recognize big‐headed grass snakes from western Transcaucasia as a distinct species, N. megalocephala. Based on phylogenetic analyses of a 1984‐bp‐long alignment of mtDNA sequences (ND4+tRNAs, cyt b) of 410 grass snakes, a nearly range‐wide phylogeography is presented for both species. Within N. natrix, 16 terminal mitochondrial clades were identified, most of which conflict with morphologically defined subspecies. These 16 clades correspond to three more inclusive clades from (i) the Iberian Peninsula plus North Africa, (ii) East Europe and Asia and (iii) West Europe including Corso‐Sardinia, the Apennine Peninsula and Sicily. Hypotheses regarding glacial refugia and postglacial range expansions are presented. Refugia were most likely located in each of the southern European peninsulas, Corso‐Sardinia, North Africa, Anatolia and the neighbouring Near and Middle East, where the greatest extant genetic diversity occurs. Multiple distinct microrefugia are inferred for continental Italy plus Sicily, the Balkan Peninsula, Anatolia and the Near and Middle East. Holocene range expansions led to the colonization of more northerly regions and the formation of secondary contact zones. Western Europe was invaded from a refuge within southern France, while Central Europe was reached by two distinct range expansions from the Balkan Peninsula. In Central Europe, there are two contact zones of three distinct mitochondrial clades, and one of these contact zones was theretofore completely unknown. Another contact zone is hypothesized for Eastern Europe, which was colonized, like north‐western Asia, from the Caucasus region. Further contact zones were identified for southern Italy, the Balkans and Transcaucasia. In agreement with previous studies using morphological characters and allozymes, there is no evidence for the distinctiveness of N. megalocephala. Therefore, N. megalocephala is synonymized with N. natrix.

Obsidian Sources in the Regions of Erzurum and Kars (North‐East Turkey): New Data

The obsidian sources on the Erzurum–Kars Plateau have not been extensively surveyed, and their geochemical signatures are still poorly understood. Yet a significant number of artefacts from archaeological sites in Georgia and Armenia have produced chemical compositions that are unrelated to any Turkish or Caucasian source analysed so far. Their origins may lie in these poorly known deposits. The objective of the collaborative project undertaken by the University of Erzurum and the French mission ‘Caucasus’ is to study the sources of obsidian in the Erzurum and Kars regions, in order to shed light on the intensity of exploitation of this material, and to highlight the exchange networks that may have existed between north‐eastern Turkey and the southern Caucasus. The analyses that we have carried out on the samples taken during this exploratory survey have enabled a definite extension of the territory of circulation of this obsidian to western Transcaucasia. The lack of knowledge concerning the diffusion of obsidian from the regions of Erzurum and Kars thus appears for the moment mainly related to insufficient geochemical characterization of the sources, confirming the importance of future surveys.

Hidden diversity in bent-winged bats (Chiroptera: Miniopteridae) of the Western Palaearctic and adjacent regions: implications for taxonomy

The taxonomic status of bent-winged bats (Miniopterus) in the Western Palaearctic and adjacent regions is unclear, particularly in some areas of the eastern Mediterranean, Middle East and Arabia. To address this, we analysed an extensive collection of museum materials from all principal parts of this distribution range, i.e. North Africa, Europe and southwest Asia, using morphological (skull) and genetic approaches (mitochondrial DNA). Linear and geometric morphometric analysis of cranial and dental characteristics, together with molecular phylogeny, suggested that Miniopterus populations comprise four separate species: (1) M. schreibersii sensu strictissimo (s.str.) – occurring in Europe, coastal Anatolia, Levant, Cyprus, western Transcaucasia, and North Africa; (2) M. pallidus – occurring in inland Anatolia, Jordan, eastern Transcaucasia, Turkmenistan, Iran and southern Afghanistan (Kandahar); (3) a Miniopterus sp. – recorded from Nangarhar province in eastern Afghanistan, which we tentatively assign to M. cf. fuliginosus; and (4) a Miniopterus sp. with Afro-tropic affinities confirmed from south-western Arabia and Ethiopia, which we tentatively name M. cf. arenarius. The latter two species are well differentiated by skull morphology, while M. pallidus possesses very similar skull morphology to M. schreibersii. The results also suggest the existence of a possible new taxon (subspecies) within M. schreibersii s.str. inhabiting the Atlas Mountains of Morocco. © 2012 The Linnean Society of London