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Citation: Mozaffarian F (2013) A preliminary study on the distribution patterns of endemic species of Fulgoromorpha (Hemiptera, Auchenorrhyncha) in Iran. In: Popov A, Grozeva S, Simov N, Tasheva E (Eds) Advances in Hemipterology. ZooKeys 319: 231–248. doi: 10.3897/zookeys.319.4159
Iran is known as the most complex and varied country in southwest Asia, in terms of geography, vegetation, climate and consequently biological diversity. The rather high number of recorded endemic species of Fulgoromorpha in Iran indicates a high potential for speciation in some areas.
In this study, in order to identify the endemic zones for Fulgoromorpha of Iran, three main biogeographic regions of the country were divided into 13 primary zones, mainly according to the distribution of published and unpublished locality records of endemic species. Using Venn diagrams and cluster analyses on the primary zones, 6 final endemic zones were recognized: Caspian zone, southern slopes of Alborz, Zagros Mountains, Kerman Mountains, Khorasan Mountains, and Baluchestan and Persian Gulf coasts. Then a similarity map was produced for endemic zones using a Multidimensional analysis (Alscal) and the differences between the positions of the same zones in the similarity and geographic maps were discussed.
Zoogeography, Iran, Fulgoromorpha
Iran is located in southwest Asia between the latitudes of 25°30' and 40° north and the longitudes of 44° and 63°30' east and has a surface area of 1, 648, 195 square kilometers. It is limited by central Asia, Caspian Sea, Caucasus Mountains and Aras River to the north, Anatolian Plateau and Mesopotamian region to the west, Persian Gulf and Oman Sea which is connected to the Indian Ocean to the south and Afghanistan and Pakistan to the east. The major part of the land of Iran is covered by the Plateau of Iran, a triangle between the Alborz Mountain Range in the north, the Zagros Mountains in the west and the Sulaiman Mountain Range in Pakistan and Afghanistan.
According to Manuel
In terms of biodiversity richness, Iran is considered as an extremely complex area and wide ranges in the extremes of altitude (below sea-level in shores of the Caspian sea to 5, 770 m of the Damavan Mt.), climate (humid and nearly jungle-like forests in the north to arid places in Dasht-e Lut with less than 100 m annual rainfall) and temperature (from -35°C in the north west to 70°C in the deserts of Dasht-e Lut) (
Biogeographic studies in Iran have been carried out mainly on plants (
Distribution of endemic Fulgoromorpha of Iran, Phytogeographical regions (Black lines) (after
The oldest zoogeographic division was performed by
A total of 235 species of Fulgoromorpha have been recorded from Iran since 1902 (
A total of 473 locality records of 117 species of planthoppers endemic to Iran were gathered, 416 of which derived from publications, 48 were newly collected and identified species in the Hayk Mirzayans Insect Museum (HMIM), and 9 were studied in the Zoological Institute of the Russian Academy of Science (ZIN). Due to the lack of the exact locality data of 3 endemic species: Phantia lactea Rusiecka, 1902 (Flatidae), Phantia putoni Rusiecka, 1902 (Flatidae) and Oliarus convergens (Melichar, 1902) (Cixiidae), they were deleted from further analyses. Then the distribution maps were prepared using Arc Map 9.3. and the country was divided into 13 primary zones according to the distribution of the endemic planthoppers, topography, climate condition and published zoogeographic zones (Fig. 1). Venn diagrams and cluster analyses were performed using Gliffy venn diagram software and NTSYS (2.02g) (
This is a small area in northern Iran, from the Caucasian-Euxino-Hyrcanian province of the Euro-Siberian region (
The district is considered as one zone (Caspian zone) in this study. Fifteen endemic planthoppers of Iran have been recorded from this zone, 9 of which are endemic to the region (Table 1 and Fig. 2). The high and wall-like Alborz Mountain separates the south coast of the Caspian Sea from the Irano-Turanian region. This barrier not only prevents the fauna from migrating to south parts but also limits the high humidity of the Caspian Sea to the north and produces two very different climatic regions in the northern and southern slopes which can be considered a major factor for differentiating the fauna of north Iran from other parts of the country. Most of the Iranian endemic species common between Hyrcanian and other regions are limited to the Alborz Mt. and are distributed in areas very close to the borders of the Hyrcanian district. Malenia masirica Dlabola, 1986 (Derbidae) is the only endemic planthopper recorded in all three main regions. There are only a few records of the presence of this species in Iran which belong mostly to the southern Zagros, the Nubo-Sindian region, close to Irano-Turan, and one record of a female specimen from the north of Iran, all made by
The number of endemic planthoppers of Iran in the main biogeographic areas.
List of endemic species of Iran, indicating endemic species of suggested endemic zones (Extracted from
Endemic species | Caspian zone | Kerman Mountains | Zagros mountains | Khorasan Mountains | Southern Slopes of Alborz | Baluchestan and Persian Gulf coasts |
---|---|---|---|---|---|---|
Family: Caliscelidae | ||||||
Adenissus baluchestanicus Dlabola, 1980 | * | |||||
Adenissus isinus Dlabola, 1980 | * | |||||
Adenissus zabolicus Dlabola, 1980 | * | |||||
Adenissus zahedanicus Dlabola, 1980 | * | |||||
Aphelonema brunneolutea Dlabola 1994 | * | |||||
Perissana dlabolai Gnezdilov & Wilson, 2006 | ||||||
Reinhardema pasagarda (Dlabola, 1982) | * | |||||
Family: Cixiidae | ||||||
Duilius v-atrum (Dlabola, 1985) | ||||||
Cixius persicus Distant, 1907 | * | |||||
Myndus genocolus Dlabola, 1985 | * | |||||
Myndus sarbazus Dlabola, 1989 | * | |||||
Anoculiarus ornatus Dlabola, 1985 | * | |||||
Eumecurus apunctatus Dlabola, 1985 | ||||||
Eumecurus baluchestanicus Dlabola, 1985 | * | |||||
Eumecurus octopus Dlabola, 1985 | * | |||||
Eumecurus superstylus Dlabola, 1985 | * | |||||
Eumecurus transpunctatus Dlabola, 1985 | * | |||||
Eumecurus vilbastei Dlabola, 1985 | * | |||||
Hyalesthes restultus Dlabola, 1994 | * | |||||
Hyalesthes zabolicus Dlabola, 1985 | ||||||
Oliarus convergens Melichar, 1902 | ||||||
Pentastira bahtiaricus (Dlabola, 1981) | * | |||||
Pentastira superspicata Dlabola, 1985 | * | |||||
Pentastira shul (Dlabola, 1985) | * | |||||
Reptalus eremicus Dlabola, 1985 | ||||||
Reptalus reductus Dlabola, 1994 | * | |||||
Reptalus ziaran Dlabola, 1985 | * | |||||
Family: Delphacidae | ||||||
Tropidocephala prasina Melichar, 1902 | * | |||||
Muirodelphax amol Dlabola, 1981 | * | |||||
Pseudaraeopus curtulus Dlabola, 1960 | * | |||||
Pseudaraeopus iranicus Dlabola, 1960 | * | |||||
Ribautodelphax hyrcanus Dlabola, 1981 | * | |||||
Family: Derbidae | ||||||
Malenia isinica Dlabola, 1986 | * | |||||
Malenia masirica Dlabola, 1986 | ||||||
Proutista jezeki Dlabola, 1981 | * | |||||
Family: Dictyopharidae | ||||||
Callodictya kazeruna (Dlabola, 1986) | * | |||||
Dictyophara albata Dlabola & Heller, 1962 | ||||||
Dictyophara exoptata Dlabola & Heller, 1962 | ||||||
Dictyophara pazukii (Dlabola, 1984) | * | |||||
Dictyophara hoberlandti Dlabola, 1974 | ||||||
Kumlika mandrita Emeljanov, 1997 | * | |||||
Nymphorgerius convergens Emeljanov, 1972 | ||||||
Nymphorgerius emeljanovi Dlabola, 1979 | * | |||||
Nymphorgerius mullah Dlabola, 1979 | * | |||||
Nymphorgerius rostratus Emeljanov, 2009 | ||||||
Family: Flatidae | ||||||
Bahuflata punctata Dlabola, 1979 | * | |||||
Derisa atratula Melichar, 1902 | ||||||
Mesophantia kanganica Dlabola, 1983 | ||||||
Mesophantia pallens Melichar, 1902 | ||||||
Mesophantia sabzevaranica Dlabola, 1983 | ||||||
Mesophantia tisina Dlabola, 1983 | ||||||
Persepolia jasmuriana Dlabola, 1982 | * | |||||
Persepolia secunda Dlabola, 1981 | * | |||||
Persepolia servadeina Dlabola, 1982 | ||||||
Phantia borazianica Dlabola, 1989 | * | |||||
Phantia crucispina Dlabola, 1989 | * | |||||
Phantia denasuta Dlabola, 1989 | * | |||||
Phantia finita Dlabola, 1989 | * | |||||
Phantia helleri Linnavuori, 1962 | ||||||
Phantia lactea Rusiecka, 1902 | ||||||
Phantia ovatospina Dlabola, 1989 | * | |||||
Phantia picea Dlabola, 1989 | * | |||||
Phantia putoni Rusiecka, 1902 | ||||||
Tisia esfandiarii Dlabola, 1981 | * | |||||
Family: Issidae | ||||||
Cavatorium ardakanum Dlabola, 1980 | * | |||||
Cavatorium bispinatum Dlabola, 1980 | * | |||||
Cavatorium quadrispinatum Dlabola, 1980 | * | |||||
Cavatorium sarbaz Dlabola, 1980 | * | |||||
Eusarima iranica Gnezdilov & Mozaffarian 2011 | * | |||||
Inflatodus astyages Dlabola, 1982 | * | |||||
Inflatodus kyaxares Dlabola, 1982 | * | |||||
Inflatodus persicus (Dlabola, 1981) | * | |||||
Inflatodus viridans (Dlabola, 1974) | * | |||||
Iranodus amygdalinus Dlabola, 1980 | ||||||
Iranodus dumetorus (Dlabola, 1981) | * | |||||
Iranodus khatunus (Dlabola, 1981) | ||||||
Iranodus nishabur Dlabola, 1982 | * | |||||
Iranodus repandus (Dlabola, 1981) | * | |||||
Iranodus transversalis Dlabola, 1980 | * | |||||
Mycterodus astragalicus Dlabola, 1974 | * | |||||
Mycterodus demavendinus Dlabola, 1981 | * | |||||
Mycterodus elbursicus (Logvinenko, 1974) | * | |||||
Mycterodus fagetophilus Dlabola, 1980 | * | |||||
Mycterodus guilanicus Dlabola, 1981 | * | |||||
Mycterodus hezarmeshedi Dlabola, 1980 | * | |||||
Mycterodus inassuetus Dlabola, 1981 | * | |||||
Mycterodus kandavanicus Dlabola, 1980 | ||||||
Mycterodus krameri Dlabola, 1974 | ||||||
Mycterodus lanceatus Dlabola 1997 | * | |||||
Mycterodus peterseni Dlabola, 1980 | * | |||||
Mycterodus sexpunctatus Dlabola, 1980 | ||||||
Mycterodus shahrudicus Dlabola, 1980 | * | |||||
Pentissus bamicus Dlabola, 1980 | ||||||
Phasmena adyoungi Dlabola, 1982 | * | |||||
Phasmena telifera Melichar, 1902 | * | |||||
Quadriva aurita (Dlabola, 1982) | * | |||||
Quadriva dehbakrina (Dlabola, 1980) | * | |||||
Quadriva lassa (Dlabola, 1981) | * | |||||
Quadriva proxima (Dlabola, 1980) | * | |||||
Quadriva sabzevarana (Dlabola, 1980) | * | |||||
Quadriva taftanica (Dlabola, 1980) | * | |||||
Quadriva tangesarhena (Dlabola, 1980) | ||||||
Scorlupaster emersum (Dlabola, 1981) | * | |||||
Family: Kinnaridae | ||||||
Perloma boroumandi (Dlabola, 1981) | * | |||||
Perloma satrapa (Dlabola, 1981) | ||||||
Perloma zarudnyi (Emeljanov, 1984) | * | |||||
Family: Meenoplidae | ||||||
Anigrus farsicus Dlabola, 1986 | * | |||||
Family: Nogodinidae | ||||||
Hadjia nerii Dlabola, 1981 | * | |||||
Hadjia quadrifasciata Dlabola, 1981 | * | |||||
Iranissus ephedrinus Dlabola, 1980 | ||||||
Morsina persica Melichar, 1902 | ||||||
Philbyella glarea Dlabola & Heller, 1962 | ||||||
Family: Ricaniidae | ||||||
Ricania soraya Dlabola, 1983 | * | |||||
Family: Tettigometridae | ||||||
Tettigometra demavenda Dlabola, 1981 | * | |||||
Family: Tropiduchidae | ||||||
Kazerunia leguaniforma Dlabola, 1977 | * | |||||
Kazerunia ochreata Dlabola, 1974 | * | |||||
Kazerunia undulata Dlabola, 1977 | * |
Historically, the north of Iran was occupied by the old Tethys 55-20 mya (Ghorbani, 2002) so the occupation should have occurred after the area got dry. During the Pliocene ice age (100, 000 years ago) the region was used as a refugee, and species spread northwards after the ice age, which may be a reason for rather small numbers of endemic species in this region.
According to
The region covers nine tenths of the land of Iran and major parts of the Iranian Plateau, from central to south of Asia. The region consists of plains, deserts and mountains which are divided into 8 primary divisions in this study: Kerman, Karkas Mt., Humid Zagros, Dry Zagros, North Khorasan Mt., South Khorasan Mt., Southern slopes of Alborz Mt. and Deserts and Plains.
The complex of the Kerman Mts located in the southeast of the Irano-Turanian region and partially in the Nubo-Sindian (Fig. 1). The mountain range is a part of the Zagros chain but far from the other mountains and surrounded by deserts. The mountain and the deserts around have been considered as a zoogeographic zone by
Venn diagrams for the number of endemic species in Kerman Mts and other areas with common species.
The Zagros mountain chain stretches from northwest of Iran to southeast near the Strait of Hormoz and is considered as the south branches of the Alpine-Himalayan orogenic belt (
In this study the Zagros chain was divided into 3 primary zones (Fig. 1): “Humid Zagros” in the northwest which consists of higher mountains and is affected by Mediterranean winds, “Dry Zagros” in the southeast with a drier climate and rather isolated mountains as the result of wider parallel valleys, and the Karkas mountain which appears to be a rather isolated mountain surrounded by Maranjab kavir and other lower lands around. Two different zones in northwest and southeast Zagros are also applied by
A total of 36 endemic planthoppers of Iran have been recorded from Zagros Mts, 19 of which are endemic to the Zagros (Table 1), 17 endemic to Dry Zagros, and 2 in common with Dry Zagros and Karkas Mt. or Humid Zagros (Fig. 4). The Venn diagram (Fig. 4) shows there are no endemic species for Humid Zagros and Karkas Mt. and the fauna of Iranian endemic planthoppers in those primary zones are mainly in common with Dry Zagros and partially with the south Alborz and Caspian zones. Therefore, those two primary divisions of the Zagros Mountains are not considered as endemic zones. Differently from the wall-like Alborz Mt., the Zagros chain comprises series of parallel ridges and valleys. The distances between mountains increases from northwest to southeast. Although the wider lower lands between the mountains in the south of Zagros (Dry Zagros) form corridors for the distribution and migration of species, mountains surrounded by those lower lands can provide a vicariant condition for speciation or at least prevent the distribution of species to other locations. The fauna of endemic planthoppers in Dry Zagros has some species in common with the Nubo-Sindian (10) and Kerman Mt. (2) which can be explained by the rather short distance between them.
Venn diagram for the number of endemic species in Zagros Mts. (Humid Zagros, Dry Zagros and Karkas Mt.) and other areas with common species.
It is believed that Iran has been connected to central Asia since the Oligocene. Then the plateau of Iran was uplifted during the Tertiary orogeny and consequently Kope Dagh (Northeast of Iran), Hindukush and Himalaya were created (
A total of 11 Iranian endemic planthoppers have been recorded from Khorasan Mts, 5 of which are endemic to this mountain range (Table 1), 3 in the south, 1 in the north and 1 in common (Fig. 5). The cluster analysis shows the similarity of the northern and southern fauna (Fig. 6). Hence the two mountains can be considered as a same zone in the distribution pattern. While south parts, which are surrounded by Kavir-e Lut and Dasht-e Kavir to the west and lowlands in Afghanistan to the east, have more endemic species and appear to be more suitable places for speciation.
Venn diagram for the number of endemic species in Khorasan Mts and common species with other areas.
Cluster diagram for the endemic areas of Fulgoromorpha in Iran.
The Alborz mountain chain represents a north branch of the Alpine-Himalayan orogenic system (
Seventeen endemic planthoppers of Iran are recorded from the south of Alborz, 12 of which endemic to the region (Table 1, Fig. 7). The southern slopes of Alborz have an absolutely different climatic condition from the northern Slopes due to the lack of corridors for receiving the humidity of the Caspian Sea. The presence of the Dasht-e Kavir desert in the south can be another factor for preventing the distribution of populations and probably speciation. There is only one common endemic planthopper between the southern slopes of Alborz and northern Khorasan whereas three species in common with the Zagros show a relationship with the western mountains.
Venn diagram for the number of endemic species in southern slopes of Alborz and common species with other areas.
Two big salt pans or Kavirs in central Iran are undrained basins and the lands surrounding them have typical features such as clayey or sandy soil and a high amount of salt on the surface. The inner lands of these salt marshes are nearly free of vegetation (
This part of the country in southern and southeastern Iran is the only region which is not considered as Palaearctic but as a part of the Oriental. A varied terminology has been used for this region.
By subtracting the endemic species of a small part of the Kerman Mountain from the Nubo-Sindian, 54 recorded endemic species of Iran, which is more than half of the total, are distributed in this region and 32 of these are endemic to the region (Table 1). The distribution of endemic Fulgoromorpha shows three rather separate groups which are recognized as three primary zones here: Baluchestan, the Strait of Hormoz and the coasts of the Persian Gulf (Fig. 1). The cluster analysis showed the similarity of the fauna of the endemic species in the Strait of Hormoz and the Persian Gulf coasts (Fig. 8) so those two primary zones can be considered as a same zone.
Cluster diagram for the primary zones of endemic Fulgoromorpha in tjr Nubo- Sindian region.
Venn diagram for the number of endemic species of the Nubo-Sindian region in three primary zones (Numbers in the parentheses are the number of endemic species in that specific primary zone)
The final 6 endemic zones (Caspian zone, South slopes of Alborz, Khorasan Mts, Zagros Mts, Kerman Mts and Baluchestan and Persian Gulf coasts) are shown in Fig. 10. The cluster analysis on six final endemic zones of Iranian planthoppers (Fig. 6) indicates Baluchestan and Perisan Gulf coasts the only region which is considered as a part of the Oriental Region, as the most different zone from the others. Then Zagros Mts, Kerman Mts, Alborz, Khorasan Mts and Caspian zones are diverging, respectively. Alscal analysis produced a map according to the similarity of endemic zones. Rotating the similarity map canvas horizontally (Fig. 11), makes it more comparable with the geographic map (Fig. 10). The Caspian zone and the Nubo-Sindian region are located in similar situations in both maps. While the Khorasan zone is closer to the Caspian zone in the similarity map rather than its geographic situation. This similarity can be justified by the possibility of specimens migrating between two zones at the end of the wall-like mountains of Alborz at the east of the Caspian zone. This is why the southern slopes of Alborz go farther in the Alscal map and confirms the efficiency of the barrier of the Alborz mountain in separating species of the two adjacent zones. Although the Zagros mountain has rather similar situations in both maps, it is farther from the Nubo-Sindian in similarity rather than its geographic situation. The reason may be the higher number of endemic species in Baluchestan rather than in the coasts of Persian Gulf and the Strait of Hormoz close to the Zagros Mountains. The situation of the Kermann Mts in the similarity map moved from its geographic situation towards the north. The rather far distance of this zone from the others, confirms the isolation of the species living there. However, deleting the doubtful record of Morsina persica, common between the Caspian zone and the Nubo-Sindian part of the Kerman Mountains may move the situation of the Kerman zone in the similarity map towards the south.
Final endemic zones and distribution of endemic Fulgoromorpha of Iran.
A similarity map of endemic zones of Fulgoromorpha of Iran resulted from Aslcal analysis.
The records of the endemic planthoppers of Iran belong to recent 110 years (
I sincerely thank Prof. A. F. Emeljanov and Dr V. M. Gnezdilov for an opportunity to study the Iranian specimens in the Zoological Institute of the Russian Academy of Sciences in St. Petersburg (Russia) and Dr M. R. Wilson, Dr P. J. Mazzoglio, Dr I. Gjonov, Dr N. Simov and anonymous reviewers for valuable suggestions.