Research Article |
Corresponding author: Sayeh Serri ( serrisayeh@gmail.com ) Academic editor: Dominique Zimmermann
© 2016 Sayeh Serri, Johannes Frisch.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Serri S, Frisch J (2016) Species diversity, chorology, and biogeography of the Steninae MacLeay, 1825 of Iran, with comparative notes on Scopaeus Erichson, 1839 (Coleoptera, Staphylinidae). Deutsche Entomologische Zeitschrift 63(1): 17-44. https://doi.org/10.3897/dez.63.5885
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The species diversity, chorology, and biogeography of the Steninae MacLeay, 1825 (Coleoptera: Staphylinidae) in Iran is described. A total of 68 species of Stenus Latreille, 1797 and one species of Dianous Leach, 1819 is recorded for this Middle Eastern country. Dianous coerulescens korgei Puthz, 2002, Stenus bicornis Puthz, 1972, S. butrintensis Smetana, 1959, S. cicindeloides Schaller, 1783, S. comma comma Le Conte, 1863, and S. hospes Erichson, 1840 are recorded for the Iranian fauna for the first time. Records of S. cordatoides Puthz, 1972, S. guttula P. Müller 1821, S. melanarius melanarius Stephens, 1833, S. planifrons planifrons Rey, 1884, S. pusillus Stephens, 1833, and S. umbricus Baudi di Selve, 1870 for Iran are, however, implausible or proved erroneous. Based on literature records and recent collecting data since 2004, the distribution of the stenine species in Iran is mapped, and their biogeographical relationships are discussed. As far as possible, the species are attributed to the commonly accepted distribution types in the Palaearctic Region after the theory of Pleistocene refuges of the arboreal biota. With the example of the Steninae and Scopaeus Erichson, 1839 (Staphylinidae: Paederinae), a cluster analysis is performed to examine the similarity of commonly accepted geographical units of Iran. The similar biogeographic pattern of the Steninae and Scopaeus reveal the high impact of Mediterranean elements on the fauna of Iran. The Hyrcanian subregion of the Caspian refuge is identified as the most significant center of origin of Stenus in Iran.
Staphylinidae , Steninae , Dianous , Stenus , distribution, biogeography, new country records, Iran
The Steninae MacLeay, 1825 constitute a monophyletic group (
Though the taxonomic diversity of the Stenus of Iran was relatively well known, the biogeography of the Iranian species still was to be investigated. Most of our knowledge on the Stenus species of Iran we owe to Puthz (e.g.
Iran is situated in the western Iranian Plateau and constitutes one of the largest countries of the Middle East. Some 47% of Iran is covered by natural grasslands, 31% by a variety of semiarid and arid environments, 14% by arable land, and 8% by woodlands (
Due to its geographic location within the Middle Eastern transitional zone of the Palaearctic, the Oriental, and the Afrotropical Regions and its diverse geology and topography, Iran shows a significant biogeographic variety. Though the country is located in the Palaearctic Region, the fauna of southern Iran is significantly influenced by Afrotropical and Oriental faunal elements. Although the leading biogeographical concepts of Iran are based on phytogeography (e.g.
A commonly accepted concept of the zoogeographical units of Iran has not yet been published.
Despite of the unique geographical features of the country, we have insufficient information about the specific zoogeographical pattern and the degree of endemism of the Iranian fauna (
In this contribution, we present the results of a research project of the first author on the diversity and biogeography of the Steninae in Iran. The aim of the study was to identify zoogeographical patterns of the stenine fauna of Iran and to compare them with the zoogeographical patterns of the strictly riparian staphylinid genus Scopaeus Erichson, 1839, research subject of the second author, to test whether they follow the same distributional patterns.
We compile the data on the distribution of the 68 Stenus species and one species of Dianous known from Iran so far, which include both previously unpublished, mostly recent records and plausible literature records, most of which were published in the last 44 years only. Finally, a cluster analysis with paired groups using Dice Similarity Coefficient was performed to gain a concept of the similarity of the Stenus fauna of defined geographical regions of Iran. It was compared with a cladogram for the Scopaeus species of Iran provided by the same method.
This study is mainly based on the results of recent field work of the authors from 2004 to 2011. The chorological data are compiled in the MS Excel 2007 supplementary file (supplementary material 1; 967 records). In this compilation, the locality labels of the specimens usually are not cited verbatim, but standardized and completed by adding the province name. Missing GPS – coordinates were subsequently taken from Tageo.com and are indicated by rectangular brackets to distinguish them from those measured at the exact collecting site.
The samples were identified by the first author and in difficult cases confirmed by Volker Puthz, the leading specialist of Steninae. Literature records were considered only if they were confirmed by examination of the reference specimens or are plausible, because they were published by well-known Stenus specialists. The dubious records of 19 species for Iran by
The specimens referred to in this contribution are stored in Hayk Mirzayans Insect Museum, Tehran, except otherwise stated. Specimens collected by A. Senglet are kept in the Muséum d’histoire naturelle, Geneva and the private collection of Volker Puthz. The collections are abbreviated as follows (in alphabetical order):
APCE = Andreas Pütz private collection, Eisenhüttenstadt;
The general information on the distribution of the species in the results section is taken from
The subgeneric classification of Stenus has been subject to controversial discussions. According to
The distribution maps were prepared using the biodiversity software BIOOFFICE and do not include dubious literature records. The distributional patterns of all members of a particular species group are combined in one map.
The biogeographical cluster analysis was carried out using NTSYS (2.02) (
The underlying geographical subdivisions mainly follow
The geographical subdivisions of Iran used in the biogeographical cluster analysis are illustrated in Fig.
Distribution of Steninae in Iran. 1. Geographical subdivision of Iran modified after
Our expeditions between 2004 and 2011 covered most of Iran. While our samples from northern and western Iran must be looked upon as quite representative for these regions, the results from the mountains of southern Khorasan are poor and preliminary, because in this region we had a short collecting chance only. In view of the niche preferences of the Steninae species, however, we don’t expect the existence of a rich Stenus fauna in the dry, salty lowlands of the coastal regions of the Persian Gulf and the Oman Sea.
In the following species chapters, we discuss the biogeography of the 69 species of Steninae known from Iran against the background of the Pleistocene glacial refuges (
Distribution of Steninae in Iran. 7. Stenus circularis species group: ○ S. planifrons robustus; 8. Stenus clavicornis species group: ○ S. caspius, ● S. p. providus; 9. Stenus comma species group: ■ S. aereus, ○ S. asiaticus, ● S. bicornis, ▲S. biguttatus, □ S. c. comma; 10. Stenus cordatus species group: ○ S. araxis, ● S. turk; 11. Stenus crassus and S. fornicatus species groups: ■ S. crassus, △ S. formicetorum, ○ S. fornicatus; 12. Stenus glacialis species group: ➕ S. armeniacus, ▲ S. hospes, ■ S. limicola, ○ S. medus, ▲S. parcior, □ S. persicus, ● S. schah.
Distribution of Steninae in Iran. 13. Stenus guttula species group: ○ S. erythrocnemus, ● S. maculiger; 14. Stenus hopffgarteni species group: ○ S. fuscicornis; 15. Stenus humilis species group: ○ S. callidus; 16. Stenus impressus and S. incanus species groups: ○ S. heinzianus, △ S. taurus; 17. Stenus latifrons and S. melanarius species groups: ▲ S. latifrons, ○ S. incrassatus, □ S. morio, ● S. peripherus; 18. Stenus melanopus species group: ○ S. atratulus, ● S. piscator.
Distribution of Steninae in Iran. 19. Stenus mendicus species group: △ S. alienigenus, ○ S. ignotus, ● S. orientis; 20. Stenus ochropus-ludyi-coarcticollis species group: □ S. barbarae, ▲ S. confrater, ▲S. darius, ●S. derwisch, △ S. guilanensis, ■S. kambyses, ● S. martensi, ○ S. ochropus, ■ S. pieperi, ➕ S. ressli, ➕S. wittmeri; 21. Stenus pallitarsis species group: ● S. arabicus, △ S. butrintensis, ○ S. claritarsis; 22. Stenus picipes species group: ○ S. ganglbaueri, ● S. picipes; 23. Stenus pusillus species group: □ S. lenkoranus, ●S. machulkai, ● S. viti; 24. Stenus similis species group: △ S. bernhauerianus, ■ S. cicindeloides, ○ S. similis.
Dianous coerulescens, widely distributed in the temperate West Palaearctic, is recorded as far east as Kazakhstan (
Dianous coerulescens cannot be assigned to any of
Stenus proprius is recorded from Greece (
Although the species was found in Greece and is expected to occur in Anatolia, its main distribution in Central Asia concurs with the Turkestanian glacial refuge. Thus, Stenus proprius can be regarded as an expansive Turkestanian faunal element.
Stenus affaber is previously published for Lebanon, Syria, Turkey, Iran, and Kazakhstan. It is collected in the eastern Zagros Mountains, northern Fars (Fig.
Stenus ater is widespread in the West Palaearctic and recorded as far east as European Russia, Azerbaijan, and North Iran, where it reaches its eastern limit of distribution in the Turkmeno-Khorasanian Mountains at about 58°06’E (Fig.
Judging from the vast distribution in the Mediterranean, we consider Stenus ater to be an expansive Holomediterranean faunal element.
Stenus hypoproditor is distributed from the northeastern Mediterranean, Southeast Europe, Turkey, and Iran as far east as Kazakhstan, Uzbekistan, and Kyrgyzstan. In Iran, it is known from the northwest (Fig.
Stenus intricatus zoufali is distributed from southeastern Central Europe and the Balkans across Anatolia, Transcaucasia, and Iran as far east as Central Asia and Afghanistan (
Stenus mongolicus is widely distributed in Asia and the Middle East from China, Mongolia, and East Siberia as far south as Indian Kashmir, Pakistan, Afghanistan, and Iran and westwards to Anatolia and Caucasia. In Iran, the species is restricted to the Elburz and Turkmeno-Khorasanian Mountains in the northeast and recorded as far south as 35°42’N in Razavi Khorasan (Fig.
Described from Bulgaria (
Due to its distribution in Southeast Europe, Anatolia, and the Caucasus region, we consider Stenus nodipes to be an expansive Pontomediterranean species.
Stenus skoraszewskyi was described from Kars, northeastern Turkey (
These few, scattered localities in the Irano-Anatolian highlands and the Elburz do not sufficiently describe the distribution pattern of Stenus skoraszewskyi, which is why we preliminarily term it as an Irano-Anatolian species.
The nominotypical subspecies of Stenus brunnipes is widely distributed in the West Palaearctic eastwards to Iran and Turkmenistan. In Iran, the subspecies seems to be restricted to the northern mountain ranges, where it was found in the eastern Elburz and the Turkmeno-Khorasanian mountains (Fig.
The Holarctic Stenus canaliculatus is widely distributed across the temperate and northern Palaearctic as far east as Russian Far East and China. In Iran, the species is confined to the temperate north (West Azarbaijan, Ardabil, Mazandaran; Fig.
Stenus canaliculatus, a faunal element of the temperate Holarctic, cannot be assigned to any of
Stenus cautus is widely distributed across the temperate and northern Palaearctic from France to the Russian Far East. The first record for Iran was published by
No biogeographical assigning is possible for this trans-Palaearctic species.
The distribution of the polytypical Stenus planifrons reaches from Germany and Switzerland over the Balkans, Anatolia, the Levant, and the Ukraine eastwards to South Russia, Caucasia, Iran, and Turkmenistan. In Iran, the species is represented by the southeastern subspecies S. p. robustus, which is distributed in Cyprus, Lebanon, Israel, Syria, Anatolia, Iraq, and northern Iran as far east as Turkmenistan.
Due to this arched areal north, west, and east of the Mesopotamian plain, the evolutionary origin of the subspecies goes back to the Syrian Pleistocene refuge. In Iran, Stenus p. robustus reaches its southern and eastern distribution limit in the southern Zagros Mountains (29°40’N) and the Turkmeno-Khorasanian mountain ranges (59°54’E) (Fig.
This cold resistant taxon was found in altitudes up to 3250 m.
Described from the Talish Mountains (
Judging from this distribution pattern, Stenus caspius is a typical expansive Caspian faunal element.
The nominotypical subspecies of Stenus providus is widespread in the West Palaearctic as far east as Kazakhstan (
Judging from the distribution in all of the arboreal Mediterranean, Stenus providus is to be attributed to the Holomediterranean distribution type.
Stenus aereus is distributed in Caucasia, the Middle East, Central Asia, and West Siberia. In Iran, it apparently is a rare species, which we did not succeed to collect. The record from West Azarbaijan published by
The Middle Eastern Stenus asiaticus is recorded for Georgia, Turkey, Syria, Iraq, Iran, and Turkmenistan. In Iran, it is the most widespread member of the S. comma species group and distributed in the South Caspian mountains and the northern Zagros Mountains (Fig.
It is difficult, to assign Stenus asiaticus to a particular distribution type judging from the total distribution. In view of the high abundance of records in the Hyrcanian zone in the northern slopes of the Elburz Mountains and the distribution in Transcaucasia, the regions which constitute the Caspian refuge, we consider S. asiaticus to be an expansive Caspian faunal element.
Described from Macedonia and Turkey (
Due to its distribution in the northeastern Mediterranean, Stenus bicornis can be regarded as a Pontomediterranean species.
Stenus biguttatus is widely distributed across the northern Palaearctic eastwards to Russian Far East and Japan (
The enormous area of distribution of Stenus biguttatus cannot be attributed to a particular Pleistocene refuge and distribution type, respectively.
The Holarctic Stenus comma is widespread across the temperate and northern Palaearctic as far east as Russian Far East, China, and Japan. In Iran, the northern species seems to be confined to the temperate northwest. Our find in West Azarbaijan constitutes the first country record of S. c. comma for Iran (Fig.
The wide distribution of this species cannot be explained by the Pleistocene refuge theory.
Stenus araxis was described from Armenia (
The distribution pattern of Stenus araxis definitely matches the Iranian glacial refuge.
Stenus turk is recorded from Azerbaijan, Georgia, Ukraine, Turkey, Iran, and Central Asia. Both Iranian representatives of the S. cordatus species group, S. araxis and S. turk, reach their southern distributional limit in Iran and occur sympatrically in the Zagros and Elburz Mountains (Fig.
Stenus crassus is widely distributed across the temperate and northern West Palaearctic eastwards to Russia, Kazakhstan, and Mongolia (
The distribution pattern of Stenus crassus cannot be explained by the theory of Pleistocene refuges.
The Holarctic Stenus formicetorum is widely distributed in the temperate and northern Palaearctic as far east as Russian Far East and as far south as Afghanistan and northwestern Iran (Fig.
The wide distribution in the northern and temperate Palaearctic cannot be attributed to a particular Pleistocene refuge.
Unlike other members of the Stenus fornicatus species group which are endemic to the eastern parts of the Palaearctic, S. fornicatus is widely distributed in the West Palaearctic including the Mediterranean and reaches East Siberia. In Iran, the species is hitherto known from the Hyrcanian zone of the northern slopes of the South Caspian mountain ranges only (Fig.
Due to the wide distribution of this species in the Mediterranean,
Stenus armeniacus was described from Yerevan, Armenia (
In view of the widely scattered localities in the Irano-Anatolian highlands we propose an Irano-Anatolian type of distribution for Stenus armeniacus.
Stenus hospes is widely distributed in the Mediterranean Region from France and Italy across the Balkans as far east as Anatolia, the Levant, and Caucasia. The recent records from the Zagros Mountains (Fig.
Judging from its distribution pattern, Stenus hospes can be regarded as a Pontomediterranean species.
Described from the Erzincan Province in the east of Turkey (
Owing to the sparse records of Stenus limicola, it is not possible to explain its Irano-Anatolian distribution pattern with de Lattin’s hypothesis.
Because descriptions of many species of the Stenus glacialis group were based on single specimens and the species never collected again,
Stenus medus, described from Armenia, Azerbaijan, and northern Iran (
Due to its distribution in Transcaucasia and its high abundance in the Hyrcanian zone in the northern slopes of the Elburz Mountains, Stenus medus is a typical expansive Caspian faunal element.
Unlike the other representatives of the Stenus glacialis species group in Iran, S. parcior is widely distributed throughout the Balkans, Anatolia, Cyprus, Lebanon, and Iran as far east as Uzbekistan. The record from Pol-e Dokhtar in the southwestern Zagros Mountains (
The distribution of Stenus parcior in the eastern Mediterranean and the Middle East corresponds to the expansive Pontomediterranean faunal element.
Stenus persicus seems to be endemic to Azerbaijan and western Iran (Fig.
Judging from the distribution pattern in the Zagros Mountains and Northwest Iran (Ardabil Province), Stenus persicus can be regarded as an Iranian faunal element.
As far as known presently, Stenus schah is endemic to Iran. The new records, which are the first after the description, revealed its wider distribution in the Zagros Mountains.
The distribution pattern in both the Zagros and Elburz Mountains (Fig.
Stenus erythrocnemus is a Middle Eastern species, which is distributed from Anatolia and Caucasia across the Iranian Plateau eastwards to Kyrgyzstan, Tadzhikistan, Afghanistan, and Pakistan. Stenus erythrocnemus has a vast distribution in the mountain ranges of Iran (Fig.
Because the majority of records originated from the highlands of Iran (
Stenus maculiger is widely distributed in the Northeast Mediterranean and eastern Europe eastwards to South Russia and Iran. The first records for Iran were published from Ghaemshahr in Mazandaran (
Judging from the distribution pattern, Stenus maculiger belongs to the expansive Pontomediterranean faunal element.
Stenus fuscicornis is widely distributed throughout most of Europe and the Mediterranean including the Maghreb countries and Turkey. The first record for Iran was published by
Stenus fuscicornis is a Holomediterranean species according to
Stenus callidus exists in the Balkans, Caucasia, and the Middle East. According to
Due to its vast distribution in the eastern Mediterranean, Anatolia (
Stenus heinzianus was described from Damash in Gilan, Iran (
Judging from this distribution pattern, Stenus heinzianus is an expansive Caspian faunal element.
Stenus taurus was described only recently from westerly facing slopes of mountain ranges close to the Turkish border in West Azarbaijan (
It is difficult to assign Stenus taurus to a particular distribution type, because its total distribution is still unknown. However, we expect it to be endemic to the Irano-Anatolian highlands.
The Stenus latifrons species group is here proposed for S. latifrons based on the deviant morphology of the paraglossae following a recommendation of V. Puthz (pers. comm., 2015).
Stenus latifrons is widely distributed across Europe from the Balkans and Italy northwards to Scandinavia. Towards the east, the species reaches Turkey, Caucasia, and Iran. A recent record from Kazakhstan (
Stenus incrassatus is widely distributed in the northern and temperate Palaearctic from Ireland to China and Russian Far East. In Iran, we collected S. incrassatus only in the Talish Mountains (Fig.
The vast range of the trans-Palaearctic species cannot be attributed to a particular Pleistocene refuge in the West Palaearctic and Middle East, respectively.
Stenus morio is a Holarctic species with a trans-Palaearctic distribution. The first records for Iran (West and East Azarbaijan) were published only recently by
The trans-Palaearctic distribution of Stenus morio can not be explained by particular Pleistocene refuges.
Stenus peripherus was described as a subspecies of S. melanarius from Turkey (
The few localities hitherto known suggest an expanded Caspian distribution for Stenus peripherus.
The West Palaearctic Stenus atratulus is widely distributed throughout Europe, the Mediterranean including the Maghreb countries Algeria and Tunesia, and Turkey eastwards to Iran, where it is most abundant in the northwest (Fig.
Due to its Mediterranean distribution centre, we classify Stenus atratulus as a Holomediterranean species following
Stenus piscator is distributed from the eastern Mediterranean (Turkey, Cyprus, Levant) across Anatolia and northern Mesopotamia as far north as Transcaucasia and eastwards to Iran and Turkmenistan. In Iran, the species has a disjunct distribution pattern (Fig.
The arched distribution pattern west, north, and east of Mesopotamia characterizes Stenus piscator as an expansive Syrian faunal element.
Described as a subspecies of Stenus mendicus from Israel (
Due to its distribution in the arboreal regions west and east of the Mesopotamian plain, we consider Stenus alienigenus as a Syrian faunal element.
Stenus ignotus was described from the Talish Mountains near Lenkoran, Azerbaijan (
Judging from the wide distribution in the northeastern Mediterranean, Stenus ignotus belongs to the Pontomediterranean type of distribution.
Stenus orientis is restricted to the Levant, Iraq, and western Iran, where it is recorded in the South Caspian mountains as far east as about 53°30’E (eastern Mazandaran) and in the Zagros Mountains southwards to 29°24’N (Central Fars) (Fig.
Due to this arched distribution around the northern Mesopotamian plain, we consider Stenus orientis as a Syrian faunal element.
Biogeographical characterization. Except for Stenus ochropus, the Iranian species of the S. ochropus – ludyi – coarcticollis species group are endemic to the Hyrcanian forest zone, the South Caspian subunit of the Caspian glacial refuge (Fig.
Stenus barbarae was described from the Talish Mountains in Gilan, Iran (
Stenus confrater is endemic to the northern Talish Mountains in Iran and Azerbaijan (Fig.
As far as known presently, Stenus darius is endemic to the northeast-facing freshwater system of the Masuleh Valley in the southern Talish Mountains at about 37°N, 49°E (Fig.
Stenus derwisch is a Hyrcanian endemic, which is comparatively widespread in the South Caspian mountains (Fig.
Stenus guilanensis was described from Lahijan, Gilan (
Stenus kambyses is presently known only from the type locality in the southernmost Talish Mountains, Gilan (Fig.
Including the new records from western Mazandaran, the first after the description, Stenus martensi is endemic to the Hyrcanian forest zone of the Elburz, where it was collected between 50°34’E (Ramsar) and 52°49’E (Alasht) in altitudes up to 1450 m (Fig.
Stenus ochropus is the most widespread member of the ochropus-ludyi-coarcticollis species group. The West Palaearctic species is distributed from Europe, except for the Iberian Peninsula, as far east as Central Asia including Anatolia, the Levant, Caucasia, and Iran, where it is recorded from the northwest as far east as Tehran and southwards to 29°09’N in Fars (Fig.
Judging from its wide distribution in the northeastern Mediterranean, we follow
Stenus pieperi was described from Alamdeh, West Mazandaran (
Stenus ressli was described from the northern foothills of the Elburz in the Caspian plain south and east of Chalus (
Stenus wittmeri, described from Minudasht, Golestan (
The Middle Eastern Stenus arabicus is recorded from Lebanon, Israel, Jordan, Syria, Turkey, and Iran as far north as Armenia. The record from China in
Due to its arched distribution in arboreal regions around northern Mesopotamia, Stenus arabicus can be considered as a Syrian faunal element.
The West Palaearctic Stenus butrintensis was previously known from western Europe as far east as Turkey. The record for Spain in
Due to its wide distribution in the northeastern Mediterranean, we adopt
Described from Lenkoran, Azerbaijan (
Stenus ganglbaueri is distributed from Italy and the Balkans across Ukraine, Anatolia, the Levant, Syria, Caucasia, and Iran eastwards to Turkmenistan. The species is widespread in northern Iran as far east as the Binalud Mountains (Fig.
Due to its distribution in the southeastern Mediterranean, we consider Stenus ganglbaueri to be a Pontomediterranean faunal element.
Stenus picipes is widespread in the temperate West Palaearctic as far east as western Russia, Anatolia, the Levant (Lebanon, Jordan, Syria), and Iran. The records from the northwestern Zagros Mountains (
The wide area of distribution of Stenus picipes can not be attributed to particular Pleistocene refugia of the arboreal, which is why no biogeographical assigning is possible for this West Palaearctic species.
Stenus lenkoranus was described as a subspecies of S. nanus Stephens, 1833 (
We propose an expansive Caspian distribution for Stenus lenkoranus.
Stenus machulkai, described from the Araxes Valley (
Judging from its existence in the northeastern Mediterranean, Stenus machulkai can be attributed to the Pontomediterranean type of distribution.
Stenus viti, endemic to the northern slopes of the South Caspian mountains and presently known from Iran only, is recorded from northern Gilan at 48°49’E (Astara) throughout Mazandaran eastwards to Golestan at 55°14’E (Ramian) (Fig.
This species is a Hyrcanian faunal element sensu
Described from Aulie Ata (Taraz) in southern Kazakhstan, the Central Asian Stenus bernhauerianus was also recorded from Kyrgyzstan and the Koppeh-Dagh (Fig.
Due to its distribution pattern, this rare species can be regarded as a Turkestanian faunal element.
The Euro-Siberian Stenus cicindeloides has a trans-Palaearctic distribution from the British Isles east to Japan. The species is here for the first time reported for Iran from Ardabil and Mazandaran in the northwest of the country (Fig.
It is not possiblt to assign the distribution pattern of Stenus cicindeloides to any of
Stenus similis is widely distributed throughout the West Palaearctic eastwards to Kazakhstan and Mongolia. The species exists in northwestern Iran and was recorded as far south as 30°44’N in the Zagros Mountains (Fig.
Judging from the existence in all of the arboreal Mediterranean, Stenus similis belongs to the Holomediterranean type of distribution.
The following Stenus species were for the first time reported for Iran by
East Azarbaijan: Arasbaran, 8.2005 (
The distribution of the widespread Holomediterranean Stenus aceris is confirmed as far east as Cyprus, Lebanon, and Turkey. The existence in northwestern Iran is doubtful.
Gilan (
The Palaearctic distribution of the Holarctic Stenus a. assequens reaches from Western Europe and the arboreal North Africa eastwards to the Russian Far East and China.
West Azarbaijan: Mahabad, 9.2003 (
Stenus binotatus is widely distributed across the Palaearctic from the British Isles to East Siberia. According to
Golestan: Kordkoy, 6.2005 (
Stenus circularis is widely distributed throughout Europe and western Asia as far east as Turkey, Caucasia, West Siberia, and Central Asia. In view of this West Palaearctic distribution pattern the above records could be correct.
East Azarbaijan: Arasbaran (
Stenus cyaneus is distributed in Lebanon, Israel, Jordan, Syria, and Turkey.
Sistan & Baluchestan (
This record was adopted from
Razavi Khorasan: Mashad, 3.2006 (
The Holarctic Stenus melanopus is widely distributed throughout the western Palaearctic eastwards to China. The cited records from Northeast Iran are dubious as well as an old record from “Perse méridionale” (
Ardabil: Ardabil, 6.2004 (
The area of distribution of the Pontomediterranean Stenus paludicola stretches over the Balkans, Anatolia, and the Levant. The existence in northwestern Iran is imaginable.
Gilan: Rasht (
The area of distribution of Stenus pallitarsis stretches over the arboreal Palaearctic eastwards to East Siberia and China. In view of the distribution in Caucasia, Turkey, and Central Asia, the existence in Iran is conceivable.
Iran (
Stenus picipennis is widely distributed in Europe and the Mediterranean, but unknown from Turkey, the Levant, and Caucasia. In view of this distribution pattern the record for Iran in the current edition of the Catalogue of Palaearctic Coleoptera far east of the area of distribution of S. picipennis appears doubtful. The underlying reference for this country record, which was not yet cited in the first edition of the catalogue (
Semnan (
The nominotypical form of Stenus planifrons has a Pontomediterranean distribution in southeastern Europe and Turkey. Considering the allopatry of subspecies, the record from Semnan is implausible, because the collecting site is situated deeply within the area of distribution of S. planifrons robustus in northwestern Iran and the South Caspian mountains (Fig.
East Azarbaijan: Arasbaran (
This species was described from Afghanistan (
Esfahan: Lenjan, 8.2000 (
Stenus pusillus is widely distributed across Europe and the arboreal North Africa (Morocco). As the species was never recorded for western Asia and the Middle East, respectively, the presence in Iran is implausible.
Fars: Kazerun, 10.2002 (
The area of distribution of the West Palaearctic Stenus solutus stretches from the British Isles as far east as Anatolia and Azerbaijan. Its existence in South Iran is doubtful.
East Azarbaijan: Kaleybar, 8.2006 (
In view of the confirmed distribution of the Pontomediterranean Stenus stigmula in Turkey and Transcaucasia, the records from northwestern Iran are plausible.
Ardabil: Meshkinshahr, 6.2007 (
Stenus subaeneus is widely distributed throughout the arboreal Mediterranean and Europe as far east as Lebanon, Syria, and Turkey. The records from northwestern Iran are questionable.
Semnan (
Stenus turbulentus has a restricted distribution in Greece, Turkey, Cyprus, and Israel. The existence of this species in North Iran is doubtful.
Khuzestan: Ahvaz, 4.2007 (
Remarks: In view of the restricted distribution of Stenus umbricus in France (Corse), Switzerland, and Italy, the record from Khuzestan is certainly based on a misidentification and here rejected.
As a result of this study, a total of 69 species of Steninae is recorded for Iran, the following of which represent new country records: Dianous coerulescens korgei, Stenus bicornis, S. butrintensis, S. cicindeloides, S. c. comma, and S. hospes.
As shown in Table
Faunal elements of the subfamily Steninae and genus Scopaeus in Iran with species numbers and percentage of species in each group.
Steninae | Scopaeus | ||||
---|---|---|---|---|---|
faunal element | species number | % | faunal element | species number | % |
Pontomediterranean | 14 | 20.29 | Iranian | 10 | 33.33 |
Hyrcanian | 11 | 15.94 | Irano-Anatolian | 5 | 16.67 |
Caspian | 7 | 10.14 | Syrian | 3 | 10 |
Holomediterranean | 6 | 8.69 | Holomediterranean | 2 | 6.67 |
Syrian | 6 | 8.69 | Pontomediterranean | 2 | 6.67 |
Trans-Palaearctic | 5 | 7.25 | Caspian | 2 | 6.67 |
Turkestanian | 5 | 7.25 | Hyrcanian | 2 | 6.67 |
Iranian | 4 | 5.80 | Trans-Palaearctic | 1 | 3.33 |
Holarctic | 4 | 5.80 | Palaeotropic | 1 | 3.33 |
Irano-Anatolian | 4 | 5.80 | South Iranian | 1 | 3.33 |
West Palaearctic | 3 | 4.35 | Sindian | 1 | 3.33 |
Palaeotropic | 0 | 0 | West Palaearctic | 0 | 0 |
Sindian | 0 | 0 | Holarctic | 0 | 0 |
South Iranian | 0 | 0 | Turkestanian | 0 | 0 |
Species number | 69 | 100 | Species number | 30 | 100 |
The results obtained from the biogeographical analysis of the distributional pattern of the subfamily Steninae in Iran are shown in Tables
Qualitative listing of Steninae species collected in Iran. See Material and methods section for abbreviations of the geographical subdivisions.
Species / Regions | NZ | NNW | C | G | SE | NKh | Z | SZ | Gh | Hm | SB | Khu | SC | SKh |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
Stenus aereus | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. affaber | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. alienigenus | 1 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 0 |
S. arabicus | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 0 |
S. araxis | 1 | 1 | 0 | 1 | 1 | 1 | 1 | 1 | 1 | 0 | 0 | 0 | 0 | 0 |
S. armeniacus | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. asiaticus | 1 | 1 | 1 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. ater | 1 | 1 | 1 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. atratulus | 1 | 1 | 1 | 1 | 0 | 1 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 0 |
S. barbarae | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. bernhauerianus | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. bicornis | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. biguttatus | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. brunnipes | 0 | 0 | 1 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. butrintensis | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. callidus | 1 | 1 | 1 | 0 | 1 | 0 | 1 | 1 | 1 | 1 | 0 | 1 | 0 | 0 |
S. canaliculatus | 0 | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. caspius | 1 | 1 | 1 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. cautus | 1 | 1 | 1 | 0 | 0 | 0 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 0 |
S. cicindeloides | 0 | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. claritarsis | 1 | 1 | 1 | 0 | 0 | 0 | 1 | 1 | 0 | 0 | 0 | 1 | 0 | 0 |
S. comma comma | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. confrater | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. crassus | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. darius | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. derwisch | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. erythrocnemus | 1 | 1 | 1 | 1 | 1 | 0 | 1 | 1 | 1 | 0 | 1 | 0 | 0 | 0 |
S. formicetorum | 1 | 1 | 1 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. fornicatus | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. fuscicornis | 0 | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. ganglbaueri | 0 | 1 | 1 | 0 | 1 | 1 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 |
S. guilanensis | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. heinzianus | 0 | 1 | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. hospes | 1 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. hypoproditor | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. ignotus | 0 | 1 | 1 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. incrassatus | 0 | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. intricatus zoufali | 1 | 1 | 1 | 1 | 0 | 1 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 0 |
S. kambyses | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. latifrons | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. lenkoranus | 0 | 1 | 1 | 0 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. limicola | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 0 |
S. machulkai | 1 | 1 | 1 | 1 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. maculiger | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. martensi | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. medus | 1 | 1 | 1 | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. mongolicus | 0 | 0 | 1 | 1 | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. morio | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. nodipes | 1 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. ochropus | 1 | 1 | 0 | 0 | 1 | 0 | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 0 |
S. orientis | 0 | 0 | 1 | 0 | 0 | 0 | 1 | 1 | 0 | 0 | 0 | 1 | 0 | 0 |
S. parcior | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. peripherus | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. persicus | 1 | 1 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. picipes | 1 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. pieperi | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. piscator | 1 | 0 | 0 | 0 | 1 | 1 | 1 | 1 | 1 | 0 | 0 | 1 | 0 | 0 |
S. planifrons robustus | 1 | 1 | 1 | 0 | 1 | 1 | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 0 |
S. proprius | 1 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. providus providus | 1 | 1 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. ressli | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. schah | 1 | 0 | 0 | 0 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. similis | 1 | 1 | 1 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. skoraszewskyi | 0 | 1 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. taurus | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. turk | 1 | 0 | 0 | 1 | 1 | 1 | 1 | 0 | 1 | 0 | 0 | 0 | 1 | 1 |
S. viti | 0 | 0 | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. wittmeri | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
Dianous coerulescens | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
Species number | 27 | 35 | 40 | 15 | 14 | 14 | 30 | 9 | 11 | 1 | 1 | 4 | 1 | 1 |
Percentage | 39.13 | 50.72 | 57.97 | 21.74 | 20.29 | 20.29 | 43.49 | 13.04 | 15.94 | 1.45 | 1.45 | 5.80 | 1.45 | 1.45 |
Number of Steninae faunal elements in each geographical unit. See Material and methods section for abbreviations of the geographical subdivisions.
NZ | NNW | C | G | SE | NKh | Z | SZ | Gh | Hm | SB | Khu | Sc | SKh | |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
Pontomediterranean | 7 | 8 | 7 | 2 | 3 | 2 | 9 | 2 | 3 | 1 | 0 | 1 | 0 | 0 |
Hyrcanian | 0 | 0 | 10 | 2 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
Caspian | 4 | 6 | 7 | 4 | 3 | 2 | 2 | 1 | 0 | 0 | 0 | 1 | 0 | 0 |
Holomediterranean | 3 | 4 | 6 | 3 | 1 | 3 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 0 |
Turkestanian | 2 | 1 | 1 | 2 | 2 | 4 | 1 | 0 | 1 | 0 | 0 | 0 | 1 | 1 |
Trans-Palaearctic | 2 | 5 | 4 | 0 | 0 | 0 | 2 | 0 | 1 | 0 | 0 | 0 | 0 | 0 |
Syrian | 3 | 1 | 2 | 0 | 2 | 2 | 6 | 4 | 2 | 0 | 0 | 2 | 0 | 0 |
Iranian | 4 | 3 | 1 | 2 | 3 | 1 | 4 | 2 | 2 | 0 | 1 | 0 | 0 | 0 |
Holarctic | 1 | 3 | 2 | 0 | 0 | 0 | 2 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
West Palaearctic | 1 | 2 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
Irano-Anatolian | 0 | 2 | 0 | 0 | 0 | 0 | 2 | 0 | 1 | 0 | 0 | 0 | 0 | 0 |
Species number | 27 | 35 | 40 | 15 | 14 | 14 | 30 | 9 | 11 | 1 | 1 | 4 | 1 | 1 |
The clade a “southern Coast + South Khorasan region “ is, however, likely to be an artifact, because the close similarity between these regions is caused by only one common species, S. turk, and not by a joint biogeographical history or habitat resemblance.
South Baluchestan, clade b, comprises S. erythrocnemus only. It is an outlier to the clusters with higher species numbers. Considering the arid character of this region, the existence of a rich Stenus fauna in South Baluchestan is not probable. Obviously, the diversity of the Steninae in Iran decreases considerably towards the eremial south and southeast owing to the increase of aridity and salinity of freshwater ecosystems.
The clade c comprises the mountains of Hormozgan, the southern Zagros Mountains in Fars, and the Khuzestan Plain with a total of nine species (13%). Southern Zagros Mountains and Khuzestan Plain show a high similarity (about 60%) with Hormozgan as an outlier with less similarity to them. It seems that appropriate habitats for Stenus species diminish towards the east and south, because the number of permanent water bodies decreases and their salinity rises.
The northern clade, d, constitutes the largest cluster of biogeographical regions of Iran. It comprises the northern, western, and central parts of the country, which reveal the least similarity with the southern clusters. It is made up of two sister clades. The first consists of Golestan and North Khorasan with 20 species and about 60% similarity. The next comprises the central and northern Zagros (37 species) with the Ghohrud Mountains and the southern Elburz as outliers with 40 species and is the sister clade to the other, which includes the Hyrcanian (Caspian) wet forest region of the Talish and the Elburz Mountains and the northwestern part of the Iranian mountain ranges with 52 species (about 75% of the Iranian Steninae). This significantly higher diversity impressively reflects the strong impact of the Pleistocene refuges proposed for these regions by
The results of the cluster analysis of the Iranian species of Scopaeus are provided in Table
Qualitative listing of Scopaeus species collected in Iran. See Material and methods section for abbreviations of the geographical subdivisions.
Species / Regions | NZ | NNW | C | G | SE | NKh | Z | SZ | Gh | Hm | SB | Khu | SC | SKh |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
Scopaeus alborzensis | 0 | 0 | 1 | 1 | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. azerbaidzhanus | 0 | 0 | 1 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. borumandi | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. cameroni | 1 | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. chalcodactylus | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 0 |
S. debilis | 1 | 1 | 0 | 1 | 1 | 1 | 1 | 1 | 1 | 0 | 0 | 0 | 0 | 1 |
S. ebneri | 1 | 1 | 0 | 0 | 1 | 0 | 1 | 1 | 1 | 0 | 0 | 0 | 0 | 0 |
S. farsensis | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 1 | 1 | 0 | 0 | 0 | 0 | 0 |
S. femursetosus | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. filiformis | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 1 | 1 | 1 | 1 | 0 |
S. gracilis | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 0 | 0 | 0 | 0 | 1 |
S. hyrcanus | 0 | 0 | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. iranensis | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. kermanensis | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 |
S. khnzoriani | 1 | 1 | 0 | 0 | 0 | 0 | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 0 |
S. kurdistanus | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. laevigatus | 0 | 1 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 |
S. mariae | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. minimus | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. mutatus | 0 | 1 | 1 | 1 | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. pakistanensis | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 |
S. persicus | 1 | 0 | 0 | 0 | 0 | 0 | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 0 |
S. qohrudensis | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 |
S. schillhammeri | 1 | 1 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. serriae | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. signifer | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 |
S. similis minor | 0 | 0 | 1 | 1 | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 |
S. stramineus | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 1 | 1 | 0 | 0 | 0 | 0 |
S. ulughdaghensis | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
S. zagrosensis | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 0 |
Species number | 12 | 11 | 9 | 8 | 8 | 8 | 11 | 10 | 8 | 2 | 2 | 1 | 1 | 4 |
Percentage | 40 | 36.67 | 30 | 26.67 | 26.67 | 26.67 | 36.67 | 33.33 | 36.67 | 6.67 | 6.67 | 3.33 | 3.33 | 13.33 |
Number of Scopaeus faunal elements in each geographical unit. See Material and methods section for abbreviations of the geographical subdivisions.
NZ | NNW | C | G | SE | NKh | Z | SZ | Gh | Hm | SB | Khu | Sc | SKh | |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
Iranian | 1 | 0 | 2 | 2 | 2 | 2 | 6 | 3 | 3 | 0 | 0 | 0 | 0 | 1 |
Irano-Anatolian | 4 | 3 | 1 | 0 | 1 | 0 | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 0 |
Syrian | 2 | 2 | 0 | 0 | 1 | 0 | 1 | 2 | 1 | 0 | 0 | 0 | 0 | 0 |
Caspian | 1 | 2 | 2 | 2 | 2 | 2 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 0 |
Hyrcanian | 0 | 0 | 2 | 2 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
Holomediterranean | 2 | 2 | 1 | 2 | 2 | 2 | 2 | 2 | 2 | 0 | 0 | 0 | 0 | 2 |
Pontomediterranean | 2 | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
Sindian | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 |
Paleotropic | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 1 | 1 | 1 | 1 | 0 |
Trans-Palaearctic | 0 | 1 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 |
South Iran and Oman | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 1 | 1 | 0 | 0 | 0 | 0 |
Species number | 12 | 11 | 9 | 8 | 8 | 8 | 11 | 10 | 8 | 2 | 2 | 1 | 1 | 4 |
The northern cluster, b + c, with 25 species the most speciose clade in the Scopaeus cladogram, is composed of two major subgroups.
The first (clade b) comprises the South Caspian and Turkmeno-Khorasanian mountain ranges. South Khorasan constitutes the outlier of the remaining geographical units of this clade. The southern Elburz is the outlier of the Caspian or Hyrcanian Region, Golestan, and North Khorasan. The two latter units reveal a significant faunal similarity (>85%), which is even higher than the considerable similarity of 60% in the Steninae.
The second subgroup (clade c) comprises Northwest Iran, the Zagros, and the Qohrud Mountains. It is made up of two sister clades with significant faunal similarity, a northern one comprising the Northwest and the adjacent North Zagros and a southern one with the Central and South Zagros and their outlier Ghohrud Mountains. The Scopaeus fauna shows a higher similarity between the northern, central, and southern parts of the Zagros compared to the Stenus fauna with a high similarity of the South Zagros and the southern regions of Khuzestan and Hormozgan.
The present study confirms previous findings (e.g.
The results reveal that the composition of the Steninae fauna in Iran is most strongly influenced by faunal elements of the Mediterranean refuge (Table
The second most frequent biogeographical element of the Iranian Steninae is the Caspian faunal element with a percentage of about 27%. Compared to the more widespread Holo-Caspian species (10.14%), which dispersed from the Caspian refuge from the Crimea to the southern shore of the Caspian Sea, however, the Hyrcanian faunal element with about 16% of the species has a greater impact on the stenine fauna of Iran. The Hyrcanian faunal elements originate from the Hyrcanian forest zone in the South Caspian plain and the northern slopes of the South Caspian mountains, the eastern subunit of the Caspian refuge. According to Frisch (2006: 262), the Hyrcanian subregion is a unique speciation center judging from the high numbers of endemics. Our results actually prove that it moreover constitutes the most significant center of origin for Stenus in Iran. The Caspian faunal elements constitute the main causative factor for the high similarity between Iranian Caucasia and the Hyrcanian region (more than 60%), and moreover strongly affect the Stenus fauna of the Golestan region and the northern Zagros Mountains. Although the Caspian Scopaeus have a considerably lower impact on the species diversity of the Iranian fauna (about 13%) compared to Stenus, some of them are relatively widespread and dispersed over geographical units of Iran adjacent to the Hyrcanian zone. Scopaeus chalcodactylus Kolenati even spread southwards to the mountains of Kerman.
Besides the Mediterranean and Caspian species, the widespread Holarctic, trans-Palaearctic, and West Palaearctic species (about 18% of the Iranian fauna) also have a great impact on the composition of the Steninae fauna of Iran, mostly in the more humid, temperate north and northwest of the country due to suitable habitats. On the contrary, the percentage of trans-Palaearctic species is much lower in the Iranian Scopaeus fauna (Table
The Syrian and Iranian refuges have their greatest influence on the Steninae fauna in the central, southern, and northern Zagros Mountains and almost the same effect on the Scopaeus fauna of these regions.
We employed the geographical units of Iran after
As the results obtained by
The distribution patterns of the stenine fauna of Iran as well as their zoogeographical relationships and degree of endemism were unknown prior to this study. With the example of these riparian rove beetles we demonstrated, that the Mediterranean Pleistocene refuge performs a huge influence on the recent fauna of this country – a result we did not expect in view of the geographical position of Iran within the Middle East and at the interface of the Palaearctic, Afrotropical, and Oriental zoogeographical regions. Our interesting results significantly contribute to a better knowledge of the biogeography of Iran and highlight the great importance of the Mediterranean for the understanding of the biogeography of the entire Middle East and beyond.
We are very grateful to the following colleagues for providing material from their institutions or private collections: N. Berti (†) (MHNP); G. Cuccodoro (
Distribution data of Steninae species from Iran.
Data type: Excel