Research Article |
Corresponding author: Ľuboš Hrivniak ( lubos.hrivniak@gmail.com ) Academic editor: Ben Price
© 2020 Ľuboš Hrivniak, Pavel Sroka, Jindřiška Bojková, Roman J. Godunko, Javid Imanpour Namin, Samereh Bagheri, Farshad Nejat, Ashgar Abdoli, Arnold H. Staniczek.
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:
Hrivniak L, Sroka P, Bojková J, Godunko RJ, Namin JI, Bagheri S, Nejat F, Abdoli A, Staniczek AH (2020) Diversity and distribution of Epeorus (Caucasiron) (Ephemeroptera, Heptageniidae) in Iran, with descriptions of three new species. ZooKeys 947: 71-102. https://doi.org/10.3897/zookeys.947.51259
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Combining morphological and molecular data in an integrative approach, three new mayfly species of Epeorus (Caucasiron) are described. These include Epeorus (Caucasiron) alborzicus Hrivniak & Sroka, sp. nov. and Epeorus (Caucasiron) shargi Hrivniak & Sroka, sp. nov. from northern Iran, and Epeorus (Caucasiron) zagrosicus Hrivniak & Sroka, sp. nov. from central Iran. They are unambiguously delimited using both distance-based and likelihood-based approaches in the analyses of barcode COI sequences. Each new species is compared with other species of the subgenus and morphological diagnostic characters are provided. Based on extensive sampling of streams throughout the country, the distribution and habitat preferences of all Caucasiron species in Iran are assessed. Altogether, there are now six species recorded, among them also E. (C.) nigripilosus Sinitshenkova, 1976 is reported for the first time in Iran. Five species are distributed in the Alborz Mts. in northern Iran, one species was found in the Zagros Mts. in central Iran.
barcoding, Caucasus, diversity, mayflies, Middle East, taxonomy
The genus Epeorus Eaton, 1881, subgenus Caucasiron Kluge, 1997 represents a group of mountainous mayflies distributed in Palaearctic region.
Caucasiron occurs in the Eastern Mediterranean (Samos and Cyprus Island), Anatolia, Caucasus, and central and western Asia (
The highest species richness of Caucasiron and a remarkable regional and local endemism was found in the Caucasus Mountains (
Individual Caucasiron species exhibit different distribution patterns within the Caucasus region varying from an endemic distribution in the Greater Caucasus to a wide distribution covering distant regions in the Pontic Mountains, Lesser Caucasus, Zagros, and Alborz Mountains (
Based on morphology and molecular analyses, we describe in this integrative study, two new species of Caucasiron from the Alborz Mountains and one new species from the Zagros Mountains. We provide morphological diagnostic characters of the three new species and differential diagnoses between all species known from the Caucasus and adjacent areas, plus an analysis of respective COI sequences. Following recent studies on Iranian mayflies by
The main objectives of this study are to (i) describe the morphology of three new Caucasiron species and provide their differential diagnoses, (ii) apply the molecular species delimitation methods using analytical tools for the single-locus COI dataset, (iii) provide basic information about habitat requirements of the new species, and (iv) summarize the distribution of all Caucasiron species recently known from Iran.
The material used for this study was collected by J. Bojková, T. Soldán, J. Imanpour Namin, and S. Bagheri in April and May 2016–2018, and A. Staniczek, M. Pallmann, R. J. Godunko, and F. Nejat in April and May 2017. All specimens were preserved in 75–96% EtOH and are deposited in the collections of the Biology Centre of the Czech Academy of Sciences, Institute of Entomology, České Budějovice, Czech Republic (
Parts of specimens were mounted on microscopic slides using HydroMatrix (MicroTech Lab, Graz, Austria) mounting medium. In order to remove the muscle tissue for an investigation of the cuticular structures, specimens were left overnight in a 10% solution of NaOH prior to slide mounting. Drawings were made using a stereomicroscope Olympus SZX7 and a microscope Olympus BX41, both equipped with a drawing tube. Photographs were obtained using Leica DFC450 camera fitted with macroscope Leica Z16 APO and folded in Helicon Focus version 5.3 X64. All photographs were subsequently enhanced with Adobe Photoshop CS5. Diagnostic characters for the description of larva were chosen according to
Total genomic DNA of the species (4–8 specimens/species) was extracted from legs using the DEP-25 DNA Extraction Kit (TopBio s.r.o., Prague, Czech Republic) according to the manufacturer’s protocol. Mitochondrial cytochrome oxidase subunit I (COI) was sequenced according to
Species were delimited using the single locus (COI) coalescence based General Mixed Yule Coalescent model (GMYC,
Inter- and intra-specific K2P pairwise genetic distances were calculated in MEGA 7 (
All of the species described below are attributed to the subgenusCaucasiron within the genus Epeorus based on the presence of projections on the costal rib of gill plates II–VII, and the presence of medio-dorsally directed hair-like setae located on the anterior margin of the head (see
Holotype : female mature larva: IRAN, Mazandaran Province, Panjab village, unnamed brook (LT of Haraz River); 36°05'52.8"N, 052°15'16.0"E (locality no. 152); 955 m a.s.l.; J. Bojková, T. Soldán, J. Imanpour Namin, S. Bagheri leg., 9.5.2018, SMNS_EPH_010056.
Paratypes : 38 female larvae (3 mounted on slide), 10 male larvae (2 mounted on slide): same data as holotype, SMNS_EPH_010056. DNA extracted from 1 female (code: IR11, stored in EtOH) and 2 males (codes: IR12 and IR14, both stored in EtOH).
33 female larvae, 24 male larvae: IRAN, Tehran Province, Zayegan village, Lalan River; 35°58'39.2"N, 051°34'56.5"E (locality no. 55); 2290 m a.s.l.; A. Staniczek, M. Pallmann, R. J. Godunko, F. Nejat leg., 8.5.2017, SMNS_EPH_007617.
1 female larva: IRAN, Golestan Province, above Chah-e Ja village, unnamed brook (RT of river flowing to Fazelabad); 36°40'22.8"N, 054°46'37.9"E (locality no. 104); 1450 m a.s.l.; J. Bojková, T. Soldán, J. Imanpour Namin leg., 27.4.2018. DNA extracted specimen (code: IR13, stored in EtOH).
17 female larvae (3 mounted on slide), 6 male larvae: IRAN, Alborz Province, 2.5 km W of Asara village, Karaj River; 36°01'52.1"N, 051°13'10.0"E (locality no. 58); 1890 m a.s.l.; A. Staniczek, M. Pallmann, F. Nejat leg., 10.5.2017, SMNS_EPH_007627.
The holotype and 50 paratypes are deposited in
8 larvae: same data as holotype, SMNS_EPH_010056; young instars or damaged specimens.
13 larvae: IRAN, Mazandaran Province, NE of Kahrud village, unnamed brook (LT of Haraz River); 36°03'42.7"N, 052°15'24.8"E (locality no. 153); 1020 m a.s.l.; J. Bojková, T. Soldán, J. Imanpour Namin, S. Bagheri leg., 9.5.2018.
2 larvae: IRAN, Mazandaran Province, 3.5 km E of Polour village, Lasem Rud (RT of Haraz River); 35°50'09.4"N, 052°04'38.4"E (locality no. 73); 2100 m a.s.l.; A. Staniczek, M. Pallmann, F. Nejat leg., 14.5.2017, SMNS_EPH_007680; 17 larvae: S. Bagheri leg., 16.4.2018.
1 larva: IRAN, Mazandaran Province, 1.5 km S of Part Kola village, Shirin Rud (LT of Sefidrud); 36°9'04.3"N, 053°20'54.7"E (locality no. 63); 750 m a.s.l.; A. Staniczek, M. Pallmann, F. Nejat leg., 11.5.2017, SMNS_EPH_007641; 10 larvae: S. Bagheri leg., 5.4.2018.
7 larvae: IRAN, Mazandaran Province, 3.5 km W of Razan village, Baladeh River; 36°11'39.6"N, 052°8'34.6"E (locality no. 73); 1360 m a.s.l.; A. Staniczek, M. Pallmann, F. Nejat leg., 14.5.2017, SMNS_EPH_007677.
1 larva: IRAN, Tehran Province, Lalan village, Lalan River; 35°59'50.3"N, 051°34'51.0"E (locality no. 53); 2438 m a.s.l.; A. Staniczek, M. Pallmann, R. J. Godunko, F. Nejat leg., 8.5.2017, SMNS_EPH_007613.
17 larvae: IRAN, Tehran Province, Igol village, Fasham River; 35°55'11.2"N, 051°28'51.3"E (locality no. 56); 2020 m a.s.l.; A. Staniczek, M. Pallmann, R. J. Godunko, F. Nejat leg., 8.5.2017, SMNS_EPH_007618.
10 larvae: IRAN, Alborz Province, 4 km NW of Shahrestanak village, Shahrestanak River; 35°59'01.2"N, 051°19'09.6"E (locality no. 57); 2100 m a.s.l.; A. Staniczek, M. Pallmann, F. Nejat leg., 10.5.2017, SMNS_EPH_007622.
The species name refers to the type locality and distribution of the species in the Alborz mountain range.
Larvae inhabit small streams (2–8 m width, 20–50 cm depth) at high altitudes (six of eleven localities at approx. 2000 m a.s.l.) in the central Alborz (Fig.
General colouration of larvae yellowish brown with dark brown maculation. Body length of mature larvae: 13.3–15.8 mm (female), 10.3–11.3 mm (male). Length of cerci approximately 1.3× body length.
Head.
Shape trapezoidal; anterior and lateral margin rounded, posterior margin rounded in female, slightly rounded or nearly straight in male (Fig.
Epeorus (Caucasiron) alborzicus sp. nov., larva: A habitus in dorsal view B habitus in ventral view C habitus in lateral view D head of male in dorsal view E head of female in dorsal view F middle leg in dorsal view G distal part of abdomen in ventral view H–J colouration of abdominal terga K–M colouration of abdominal sterna.
Mouthparts. Labrum (Fig.
Epeorus (Caucasiron) alborzicus sp. nov., larva: A labrum (right half in dorsal view, left half in ventral view) B incisors of right mandible in ventral view C incisors of left mandible in ventral view (both flattened on slide) D setae on dorsal surface of femora E surface and posterior margin of abdominal tergum VII F gill I G gill III H abdominal segments VIII–X I sternum IX, female J sternum IX, male K gill VII (flattened on slide) L–M gill VII (in natural position from ventral view), variability in shape.
Thorax. Pronotum anteriorly narrowed, lateral edges nearly straight. Metanotum with slight postero-medial projection. Dorsal surface covered with fine, hair-like setae (as on abdominal terga and head); sparse longer, hair-like setae along pro-, meso- and metanotal suture.
Legs. Colour pattern of femora as in Fig.
Abdominal terga. Colour pattern of abdominal terga (Fig.
Lateral margins with oblique maculae on terga I–IX, sometimes dorso-posteriorly extended. Pair of sigilla sometimes coloured, in form of short stripes or spots located antero-laterally to medial macula. Denticles on posterior margin on terga of various size, irregular and pointed (Fig.
Abdominal sterna. Yellowish, with distinct colour pattern in form of medial circular macula (Fig.
Gills. Dorsal surface of gill plate I yellowish; of gill plates II–VII greyish on anterior half, brownish (sometimes reddish) on posterior half. Ventral margin of all gill plates yellowish. Projection of gill plate III well developed (Fig.
Cerci. Yellowish brown, basally darkened.
Unknown.
The main larval diagnostic characters of E. (C.) alborzicus sp. nov. are as follows: (i) colour pattern of abdominal terga (Fig.
The combination of diagnostic characters mentioned above clearly distinguish larvae of E. (C.) alborzicus sp. nov. from all other Caucasiron species known so far. However, some of the diagnostic characters occur also in other Caucasiron species distributed in the Caucasus. The colour pattern of abdominal sterna in E. (C.) alborzicus sp. nov. is similar in E. (C.) bicolliculatus (
Epeorus (C.) alpestris differs by the characteristic colour pattern of abdominal terga (
The presence of postero-lateral projections on the abdominal tergum X is characteristic for two species distributed in the Caucasus, E. (C.) magnus, E. (C.) nigripilosus, and sporadically also in E. (C.) znojkoi. Epeorus (C.) magnus differs from E. (C.) alborzicus sp. nov. in the absence of colouration of abdominal sterna and the characteristic setation on the dorsal margin of labrum (numerous thickened bristle-like setae, Hrivniak et al. in prep.). Epeorus (C.) nigripilosus can be separated from E. (C.) alborzicus sp. nov. by the presence of the distinct medial hypodermal femur spot and unique colour pattern of abdominal sterna (
Two species, E. (C.) soldani and E. (C.) sinitshenkovae, are lacking a medial hypodermal femur spot just like E. (C.) alborzicus sp. nov. Both can be separated from the latter by the absence of postero-lateral projections on tergum X, narrower gill plates VII (in natural position from ventral view), and the absence of a distinct colour pattern of abdominal sterna. Additionally, E. (C.) soldani differs from E. (C.) alborzicus sp. nov. by the presence of flattened setae on the surface of abdominal terga (
Other Caucasiron species distributed in the Caucasus and adjacent areas do not share important diagnostic characters with E. (C.) alborzicus sp. nov. All of these species can be easily distinguished by the following combination of characters: (i) absence of the colour pattern of abdominal sterna and presence of the medial hypodermal femur spot in E. (C.) turcicus, E. (C.) longimaculatus, E. (C.) shargi sp. nov. and (ii) colour pattern of abdominal terga and sterna in E. (C.) caucasicus (
Epeorus (Caucasiron) shargi sp. nov., larva: A habitus in dorsal view B habitus in ventral view C habitus in lateral view D head of male in dorsal view E head of female in dorsal view F middle leg in dorsal view G distal part of abdomen in ventral view H–K colouration of abdominal terga L colouration of abdominal sterna.
Holotype : female mature larva: IRAN, Golestan Province, Shirinabad village, unnamed river; 36°48'01.4"N, 055°01'05.8"E (locality no. 108); 740 m a.s.l.; J. Bojková, T. Soldán, J. Imanpour Namin leg., 27.4.2018, SMNS_EPH_010057.
Paratypes :
19 female, 11 male larvae: same data as holotype.
36 female (5 mounted on slide), 25 male (1 mounted on slide) larvae: IRAN, Golestan Province, above Chah-e Ja village, unnamed brook (RT of river flowing to Fazelabad); 36°40'22.8"N, 054°46'37.9"E (locality no. 104); 1450 m a.s.l.; J. Bojková, T. Soldán, J. Imanpour Namin leg., 27.4.2018. DNA extracted from 2 females (codes: IR23 and IR24, mounted on slides).
19 female (3 mounted on slide), 7 male (1 mounted on slide) larvae: IRAN, Golestan Province, below Chah-e Ja village (main valley), unnamed river flowing to Fazelabad, 36°41'46.3"N, 054°47'35.0"E (locality no. 105); 1240 m a.s.l.; J. Bojková, T. Soldán, J. Imanpour Namin leg., 27.4.2018. DNA extracted from 1 female (code: IR21, mounted on slide) and 1 male (code: IR22, stored in EtOH).
The holotype (SMNS_EPH_010057) and 50 paratypes (SMNS_EPH_010057) are deposited in
3 larvae: same data as holotype; young instars or damaged specimens.
The species name derives from shargi (یقرش), which means eastern in Farsi. It refers to the distributional range of the species in the eastern part of the Alborz mountain range.
Larvae were found in three clear streams at middle altitude (740–1450 m a.s.l.) in the eastern Alborz (Fig.
General colouration of larvae yellowish brown with dark brown maculation. Body length of mature larvae 13.7–15.6 mm (female), 11.7–13.0 mm (male). Length of cerci approximately 1.1× body length.
Head.
Shape trapezoidal; anterior and lateral margin rounded, posterior margin rounded in female, slightly rounded in male (Fig.
Mouthparts. Labrum (Fig.
Epeorus (Caucasiron) shargi sp. nov., larva: A labrum (right half in dorsal view, left half in ventral view) B incisors of right mandible in ventral view C incisors of left mandible in ventral view (both flattened on slide) D setae on dorsal surface of femora E surface and posterior margin of abdominal tergum VII F gill I G gill III H abdominal segments VIII–X I sternum IX, female J sternum IX, male K gill VII (flattened on slide) L gill VII (in natural position from ventral view).
Thorax. Pronotum anteriorly narrowed, lateral edges nearly straight or slightly rounded. Metanotum with slight, blunt, postero-medial projection. Dorsal surface covered with fine hair-like setae (as on abdominal terga and head); sparse longer hair-like setae along pro-, meso- and metanotal suture.
Legs.
Colour pattern of femora as in Fig.
Abdominal terga.
Colour pattern of abdominal terga (Fig.
Abdominal sterna.
Yellowish, without distinct colour pattern. Nerve ganglia often dark brown pigmented (Fig.
Gills.
Dorsal surface of gill plate I yellowish, of gill plates II–VII greyish on anterior half and brownish to reddish on posterior half. Ventral margin of all gill plates yellowish. Projection of gill plate III well developed (Fig.
Cerci. Brownish, basally darkened.
Unknown
The main larval diagnostic characters of E. (C.) shargi sp. nov. are as follows: (i) colour pattern of abdominal terga (Fig.
Based on the colour pattern of abdominal terga and sterna, E. (C.) shargi sp. nov. resembles several species distributed in the Caucasus and adjacent areas. At first glance, E. (C.) soldani and E. (C.) turcicus are most similar. Larvae of E. (C.) soldani possess triangular maculae on abdominal terga (
Epeorus (C.) turcicus shares with E. (C.) shargi sp. nov. the lack of colouration on abdominal sterna (
Similar to E. (C.) shargi sp. nov., there is no colour pattern of abdominal sterna in several other species, namely E. (C.) longimaculatus, E. (C.) sinitshenkovae, and E. (C.) magnus. Epeorus (C.) longimaculatus can be clearly separated from E. (C.) shargi sp. nov. by (i) a distinctly narrower gill plate VII (in natural position of ventral view), (ii) flattened setae on the surface of abdominal terga (
Epeorus (C.) sinitshenkovae can be distinguished from E. (C.) shargi sp. nov. by the absence of a medial hypodermal femur spot, the characteristic colour pattern of femora (Braasch and Zimmerman 1979: 106, fig. 10), and the colour pattern of abdominal terga (
Epeorus (C.) magnus can be reliably distinguished by the presence of distinct postero-lateral projections on abdominal tergum X and characteristic setation of labrum (numerous thickened bristle-like setae, Hrivniak et al. in prep.).
All other species distributed in the Caucasus and adjacent areas differ from E. (C.) shargi sp. nov. by the distinct colour pattern of abdominal sterna, namely E. (C.) bicolliculatus (
Holotype : female larva: IRAN, Lorestan Province, 4.5 km SW of Varayeneh village, Sarab-e Gamasiab River, 34°2'46.2"N, 048°22'32.6"E (locality no. 9); 1842 m a.s.l.; A. Staniczek, M. Pallmann, A. Abdoli, F. Nejat leg., 25.4.2017, SMNS_EPH_007520.
Paratypes : 79 female larvae, 68 male larvae: same data as holotype, SMNS_EPH_007520. 6 female (2 mounted on slide), 5 male (2 mounted on slide) larvae: IRAN, Chaharmahal and Bakhtiari Province, Dimeh village, Chehme-Dimeh River, 32°30'11.6"N, 050°13'04.5"E (locality no. 45) ; 2220 m a.s.l.; A. Staniczek, M. Pallmann, R. J. Godunko, F. Nejat leg., 5.5.2017, SMNS_EPH_007707. DNA extracted from 3 females (code: IR32, stored in EtOH; codes: IR34 and IR35, mounted on slides) and 2 males (codes: IR33b and IR36, mounted on slides).
15 female (3 mounted on slide), 5 male larvae: IRAN, Kohgiluyeh and Boyer-Ahmad Province, 4 km E of Yasuj, Yasuj fall, 30°40'34.7"N, 051°37'35.6"E (locality no. 37); 2060 m a.s.l.; A. Staniczek, M. Pallmann, R. J. Godunko, F. Nejat leg., 4.5.2017, SMNS_EPH_007568. DNA extracted from 2 females (code: SP38, mounted on slide; code: IR33a, stored in EtOH) and 1 male (code: SP37, stored in EtOH).
2 female, 2 male larvae: IRAN, Chaharmahal and Bakhtiari Province, 5 km W of Chelgerd, Kouhrang River, 32°28'9.3"N, 050°5'26.2"E (locality no. 46); 2402 m a.s.l.; A. Staniczek, M. Pallmann, R. J. Godunko, F. Nejat leg., 5.5.2017, SMNS_EPH_007689.
The holotype and 100 paratypes are deposited in
42 larvae: same data as holotype; young instars or damaged specimens.
1 male larva: IRAN, Chaharmahal and Bakhtiari Province, 4 km E. of Bajgiran, Dehno River, 31°54'26.2"N, 050°42'20.6"E (locality no. 50); 1721 m a.s.l.; A. Staniczek, M. Pallmann, R. J. Godunko, F. Nejat leg., 6.5.2017, SMNS_EPH_007606.
The species name refers to its known records in the Zagros mountain range.
Larvae were found in five streams of different size at high altitude, above 1700 m a.s.l. Three streams were strongly turbulent rivers with very coarse bed substrate flowing in high-mountain valleys (Fig.
General colouration of larvae yellowish brown with dark brown maculation. Body length of mature larvae 13.5–14.5 mm (female), 10.0–11.0 mm (male). Length of cerci approximately 1.3× body length.
Head.
Shape trapezoidal; anterior and lateral margin rounded, posterior margin slightly rounded or nearly straight (Fig.
Epeorus (Caucasiron) zagrosicus sp. nov., larva: A habitus in dorsal view B habitus in ventral view C habitus in lateral view D head of male in dorsal view E head of female in dorsal view F middle leg in dorsal view G distal part of abdomen in ventral view H–J colouration of abdominal terga K colouration of abdominal sterna.
Head dimensions of mature larvae: length 2.6–2.7 mm, width 3.6–4.0 mm (female); length 2.3–2.4 mm, width 3.3 mm (male). Head width/length ratio: 1.36–1.49 (both male and female).
Dorso-medial part with indistinct brown rectangular or oval macula, sometimes reduced to pair of stripes. Rounded maculae under lateral ocelli and blurred or triangular maculae near inner edges of compound eyes. Pair of pale stripes extending from lateral ocelli to lateral edges of head. Pair of maculae located along coronal suture. Compound eyes dark grey to black in female, brownish and basally blackish in male mature larva. Ocelli dark grey to black, basally paler. Antennae yellowish-brown, scapus and pedicellus darkened. Anterior margin densely covered with hair-like setae extending to lateral margins and directed medio-dorsally. Dorsal surface covered with fine hair-like setae and sparsely distributed stick-like setae. Sparse longer, fine, hair-like setae located posteriorly to eyes.
Mouthparts.
Labrum (Fig.
Epeorus (Caucasiron) zagrosicus sp. nov., larva: A labrum (right half in dorsal view, left half in ventral view) B incisors of right mandible in ventral view C incisors of left mandible in ventral view (both flattened on slide) D setae on dorsal surface of femora E surface and posterior margin of abdominal tergum VII F gill I G gill III H–I abdominal segments VIII–X J sternum IX, female K sternum IX, male L gill VII (flattened on slide) M–O gill VII (in natural position from ventral view), variability in shape.
Thorax. Pronotum anteriorly narrowed, lateral edges nearly straight. Metanotum with slight postero-medial projection. Dorsal surface covered with fine hair-like setae (as on abdominal terga and head); sparse longer hair-like setae along pro, meso- and metanotal suture.
Legs.
Colour pattern of femora as in Fig.
Results of the molecular species delimitation: A distribution of K2P pairwise genetic distances B mean pairwise genetic distances between new species and all Caucasian Caucasiron species known C COI gene tree with the results of molecular species delimitation analyses and morphology (node supports for species are indicated).
Abdominal terga.
Colour pattern of abdominal terga includes transversal stripe along anterior margin of terga I–IX (X) medially extending to triangular, short rectangular or stripe-like medial macula on terga (III) IV–IX (transversal stripe sometimes not distinctly extended, for variability see Fig.
Abdominal sterna.
Yellowish, with distinct colouration pattern consisting of anteriorly widened pair of stripes (medio-anterior sigilla) on terga II–VIII (Fig.
The map showing the occurrence of Epeorus (Caucasiron) spp. at all localities investigated in Iran. Colour of symbols shows the occurrence of species: green – Epeorus (Caucasiron) zagrosicus sp. nov., dark blue – Epeorus (Epeorus) zaitzevi, and violet – all other Epeorus (Caucasiron) species. Black symbols show collection points where no species of Epeorus was found. The letter H shows the locality of the respective holotype.
Gills.
Dorsal surface of gill plate I yellowish; of gill plates II–VII greyish on anterior half and brownish to reddish on posterior half. Ventral margin of all gill plates yellowish. Projection of gill plate III well developed (Fig.
Cerci. Brownish, basally darkened.
Unknown
The main larval diagnostic characters of E. (C.) zagrosicus sp. nov. are as follows: (i) colour pattern of abdominal sterna (Fig.
Distribution of Epeorus (Caucasiron) species in northern Iran. Colour of symbols shows the occurrence of species: red – E. (C.) alborzicus sp. nov., light blue – E. (C.) shargi sp. nov., violet – E. (C.) cf. znojkoi, black – E. (C.) iranicus, yellow – E. (C.) nigripilosus. The letter H shows the localities of holotypes. Black symbols with white cross show unrevised records of E. (C.) iranicus.
Based on the colour pattern of abdominal sterna, E. (C.) zagrosicus sp. nov. is most similar to E. (C.) caucasicus and E. (C.) iranicus. Both latter species possess pigmented medio-anterior sigilla forming a pair of oblique stripes on abdominal sterna II–VIII (e.g.,
Distinct postero-lateral projections on the tergum X are characteristic for E. (C.) magnus, E. (C.) nigripilosus, and E. (C.) alborzicus sp. nov. Small projections are also sporadically present in E. (C.) znojkoi. E. (C.) magnus can be easily distinguished from E. (C.) zagrosicus sp. nov. by the absence of colour pattern of abdominal sterna, the absence of a medial hypodermal femur spot, and setation on dorsal margin of labrum (numerous thickened bristle-like setae, Hrivniak et al., in prep.). E. (C.) nigripilosus and E. (C.) alborzicus sp. nov. differ by a typical colouration pattern of abdominal sterna (
Photos of the localities of the new Epeorus (Caucasiron) species described herein: A unnamed brook near Panjab village – type locality of E. (C.) alborzicus sp. nov. B unnamed brook near Kahrud-e Bala village – locality of E. (C.) alborzicus sp. nov. C unnamed river near Shirinabad village – type locality of E. (C.) shargi sp. nov. D unnamed brook near Chah-e Ja village – locality of E. (C.) shargi sp. nov. E Gamasiab River near Varayeneh village – type locality of E. (C.) zagrosicus sp. nov. F Yasuj fall near Yasuj village – locality of E. (C.) zagrosicus sp. nov.
The presence of a medial hypodermal femur spot makes E. (C.) zagrosicus sp. nov. slightly similar to E. (C.) turcicus and E. (C.) alborzicus sp. nov. However, the presence of the characteristic pattern of abdominal sterna in E. (C.) alborzicus sp. nov. (Fig.
The other five species distributed in the Caucasus, namely E. (C.) sinitshenkovae, E. (C.) alpestris, E. (C.) bicolliculatus, E. (C.) longimaculatus, and E. (C.) soldani, do not share any important diagnostic characters with E. (C.) zagrosicus sp. nov. Nevertheless, E. (C.) sinitshenkovae and E. (C.) alpestris can be separated from E. (C.) zagrosicus sp. nov. by the absence of a medial hypodermal femur spot, overall colouration of the dorsal surface of femora (E. (C.) sinitshenkovae, Braasch and Zimmerman 1979: 106, fig. 10), and the different colouration of abdominal sterna (E. (C.) alpestris,
The GMYC model provided significantly better fit to COI gene tree than the null model expecting uniform coalescent branching rates across entire tree (likelihood ratio test = 3.671927e-06***). The GMYC estimated 15 species (CI=13–19) consisting of 14 ML clusters and one singleton (CI = 12–16). All three newly described species were confirmed, and the overall number of delimited GMYC species corresponded well to morphologically defined species within Caucasiron (Fig.
The ABGD analysis of the COI distance matrix recognized 15 stable groups within initial partition. All groups corresponded well to morphologically defined species and were congruent with GMYC analysis. All three newly proposed species were recognized as distinct groups (Fig.
Specimens of the genus Epeorus were found in 68 localities of all 254 localities investigated by us in 2016–2018 (Fig.
List of records of the Caucasiron species found in Iran (three new species are not included). Abbreviations: RT – right tributary; LT – left tributary; JB – Jindřiška Bojková; TS – Tomáš Soldán; IN – Javid Imanpour Namin; SB – Samereh Bagheri; AHS – Arnold H. Staniczek; MP – Milan Pallmann; RJG – Roman J. Godunko; FN – Farshad Nejat; AA – Ashgar Abdoli; HV – H. Valikhani; PT – P. Taban. Number of specimens includes larvae.
Species | Province | Stream | Locality | Nearest settlement | Altitude | Latitude (N) / Longitude (E) | Sampling date | Collector/reference | Number of specimens |
---|---|---|---|---|---|---|---|---|---|
E. (C.) iranicus | Alborz | Karaj R., Shahrestanak branch | SE of Shahrestanak | Chavar Chalun | 2220 | 35°57'45.8"N, 051°21'59.7"E | 1.9.2016 | AA, HV, PT | 4 |
Ardabil | unnamed brook | in Alvaresi (below Alvares ski area) | Sarein | 2235 | 38°09'38.0"N, 047°56'21.0"E | 17.5.2016 | JB, TS, IN; |
2 | |
Mazandaran | Koshk Sara R. | in Kosh Sara | Chalus | 18 | 36°37'57.7"N, 051°28'04.4"E | 25.9.2013 |
|
2 | |
Mazandaran | Firuz Abad R. | near Dasht Nazir | Marzan Abad | 929 | 36°24'33.8"N, 051°24'42.5"E | 11.9.2014 |
|
80 | |
Mazandaran | Haraz R. | in Gazanak | Gazanak | 1590 | 35°54'08.3"N, 052°13'30.0"E | 10.7.2013 |
|
8 | |
Mazandaran | Dalir R. | above Dalir | Marzan Abad | 2126 | 36°19'23.2"N, 051°04'27.5"E | 26.7.2014 |
|
174 | |
Mazandaran | Lasem R. | E of Polour | Polour | 2180 | 35°50'04.1"N, 052°05'07.6"E | 14.5.2017 | AHS, MP, FN | 1 | |
Mazandaran | RT of Sardab Rud | SW of Kelardasht | Kelardasht | 2020 | 36°26'06.5"N, 051°03'52.6"E | 8.3.2018 | SB | 85 | |
Mazandaran | LT of Sardab Rud | NW of Vandarbon | Kelardasht | 2250 | 36°25'53.7"N, 051°01'59.1"E | 9.3.2018 | SB | 63 | |
Mazandaran | Sardab Rud | S of Vandarbon | Kelardasht | 2290 | 36°25'23.0"N, 051°02'12.4"E | 10.3.2018 | SB | 5 | |
Tehran | Darban valley (type locality) | N of Tehran | Tehran | 2100 | 35°50'24.0"N, 051°25'19.9"E | 18.7.1970 |
|
14 | |
Tehran | Lalan R. | above Zayegan | Fasham | 2290 | 35°58'39.2"N, 051°34'56.5"E | 8.5.2017 | AHS, MP, RJG, FN | 158 | |
Tehran | Lalan R. | in Lalan | Lalan | 2440 | 35°59'50.3"N, 051°34'51.0"E | 8.5.2017 | AHS, MP, RJG, FN | 87 | |
Tehran | Ahar R. | near Igol | Fasham | 2020 | 35°55'11.2"N, 051°28'51.3"E | 8.5.2017 | AHS, MP, RJG, FN | 1 | |
Tehran | Shahrestanak R. | NW of Shahrestanak | Asara | 2100 | 35°59'01.2"N, 051°19'09.6"E | 10.5.2017 | AHS, MP, FN | 1 | |
E. (C.) nigripilosus | Alborz | Kordan R. | N of Kordan | Kordan | 1430 | 35°57'15.6"N, 050°50'25.3"E | 10.5.2017 | AHS, MP, FN | 4 |
Mazandaran | RT of Dohezar R. | N of Holu Kaleh | Tonkaboon | 880 | 36°37'37.5"N, 050°44'30.2"E | 16.6.2018 | SB | 1 | |
Mazandaran | Dohezar R. | SW of Parde Sar | Tonkaboon | 450 | 36°40'07.0"N, 050°49'20.0"E | 16.6.2018 | SB | 1 | |
Tehran | Ahar R. | near Igol | Fasham | 2020 | 35°55'11.2"N, 051°28'51.3"E | 8.5.2017 | AHS, MP, RJG, FN | 5 | |
Tehran | Shahrestanak R. | NW of Shahrestanak | Asara | 2100 | 35°59'01.2"N, 051°19'09.6"E | 10.5.2017 | AHS, MP, FN | 1 | |
E. (C.) cf. znojkoi | Alborz | Kordan R. | N of Kordan | Kordan | 1430 | 35°57'15.6"N, 050°50'25.3"E | 10.5.2017 | AHS, MP, FN | 4 |
Gilan | RT of Khara Rud | S of Paein Khara Rud (S of Pashaki) | Sangar | 210 | 37°02'29.0"N, 049°47'52.0"E | 12.5.2016 | JB, TS, IN | 71 | |
Gilan | left fork of Khara Rud | in Madarsara (S of Pashaki) | Sangar | 105 | 37°04'12.0"N, 049°46'36.0"E | 12.5.2016 | JB, TS, IN | 27 | |
Gilan | right fork of Khara Rud | in Golestansara (S of Pashaki) | Sangar | 201 | 37°02'20.0"N, 049°47'27.0"E | 12.5.2016 | JB, TS, IN | 2 | |
Gilan | Zilaki River (RT of Sefid Rud) | in Mush Bijar (E of Shahr-e Bijar) | Shahr-e Bijar | 120 | 37°00'28.0"N, 049°40'24.0"E | 13.5.2016 | JB, TS, IN | 1 | |
Gilan | Sefidab (RT of Siah Rud) | in Divarsh (NE of Shirkuh) | Tutkabon | 280 | 36°53'59.0"N, 049°35'06.0"E | 13.5.2016 | JB, TS, IN | 149 | |
Gilan | Sangdeh (LT of Shafa Rud) | W of Punel | Punel | 240 | 37°31'47.0"N, 049°00'52.0"E | 15.5.2016 | JB, TS, IN | 10 | |
Gilan | Shafa Rud | W of Punel | Punel | 240 | 37°31'47.0"N, 049°00'52.0"E | 15.5.2016 | JB, TS, IN | 6 | |
Gilan | LT of Shafa Rud | NW of Sangdeh | Sangdeh | 1345 | 37°31'46.0"N, 048°45'19.0"E | 15.5.2016 | JB, TS, IN | 50 | |
Gilan | Shakhzar R. | NE of Fuman | Fuman | 6 | 37°14'13.0"N, 049°20'43.0"E | 15.5.2016 | JB, TS, IN | 1 | |
Gilan | LT of Bala Rud | S of Siahkal | Siahkal | 490 | 37°00'31.0"N, 049°51'51.0"E | 16.5.2016 | JB, TS, IN | 16 | |
Gilan | Lunak waterfalls | S of Siahkal | Siahkal | 510 | 37°00'31.0"N, 049°51'49.0"E | 16.5.2016 | JB, TS, IN | 5 | |
Gilan | Shamrud (RT of Sefid Rud) | S of Tushi (S of Siahkal) | Siahkal | 315 | 37°03'00.0"N, 049°53'54.0"E | 16.5.2016 | JB, TS, IN | 37 | |
Gilan | Chelavand R. | W of Chelvand | Lavandvil | -4 | 38°17'20.0"N, 048°51'35.0"E | 19.5.2016 | JB, TS, IN | 2 | |
Gilan | unnamed brook | N of Chaldasht | Amlash | 1255 | 36°59'33.0"N, 050°05'19.0"E | 21.5.2016 | JB, TS, IN | 3 | |
Gilan | RT of Shalman Rud | in Bolurdekan | Amlash | 345 | 37°01'09.0"N, 050°03'51.0"E | 21.5.2016 | JB, TS, IN | 2 | |
Gilan | LT of Ghale Rudkhan | NE of Masuleh | Fuman | 885 | 37°09'47.0"N, 049°00'17.0"E | 22.5.2016 | JB, TS, IN | 66 | |
Gilan | RT of Ghale Rudkhan | NE of Masuleh | Fuman | 705 | 37°09'42.0"N, 049°01'17.0"E | 22.5.2016 | JB, TS, IN | 6 | |
Gilan | Ghale Rudkhan R. | E of Masuleh | Fuman | 370 | 37°10'02.0"N, 049°05'03.0"E | 22.5.2016 | JB, TS, IN | 4 | |
Golestan | unnamed river | in Shirinabad | Aliabad-e Katul | 740 | 36°48'01.0"N, 055°01'05.0"E | 27.4.2018 | JB, TS, IN | 50 | |
Golestan | Shirabad waterfalls | above Shirabad | Shirabad | 140 | 36°57'33.0"N, 055°01'57.0"E | 28.4.2018 | JB, TS, IN | 48 | |
Golestan | RT of Madarsu R. | E of Tangrah | Tangrah | 495 | 37°23'27.0"N, 055°48'51.0"E | 30.4.2018 | JB, TS, IN | 1 | |
E. (C.) cf. znojkoi | Mazandaran | Shirinrud | S of Part Kola | Farim | 770 | 36°09'02.5"N, 053°20'58.1"E | 11.5.2017 | AHS, MP, FN | 203 |
Mazandaran | trib. Kashpel R. | SW of Chamestan | Chamestan | 400 | 36°25'31.4"N, 052°03'38.4"E | 13.5.2017 | AHS, MP, FN | 17 | |
Mazandaran | Chelav R. | N of Pasha Kola | Pasha Kola | 820 | 36°12'24.7"N, 052°25'51.9"E | 14.5.2017 | AHS, MP, FN | 63 | |
Mazandaran | Chelav R. | NW of Pasha Kola | Pasha Kola | 570 | 36°13'28.9"N, 052°23'35.4"E | 14.5.2017 | AHS, MP, FN | 2 | |
Mazandaran | Baladeh R. | W of Razan | Razan | 1360 | 36°11'39.6"N, 052°08'34.6"E | 14.5.2017 | AHS, MP, FN | 9 | |
Mazandaran | Chai Bagh R. | E of Andar Koli | Ghaem Shahr | 200 | 36°20'30.0"N, 052°54'03.0"E | 8.5.2018 | JB, TS, IN, SB | 27 | |
Mazandaran | RT of Haraz R. | NW of Pasha Kola | Amol | 570 | 36°13'27.0"N, 052°23'36.0"E | 9.5.2018 | JB, TS, IN, SB | 5 | |
Mazandaran | LT of Haraz R. | in Panjab | Amol | 955 | 36°05'52.0"N, 052°15'15.0"E | 9.5.2018 | JB, TS, IN, SB | 1 | |
Mazandaran | unnamed brook | above Darab Kola | Neka | 135 | 36°33'10.0"N, 053°15'32.0"E | 10.5.2018 | JB, TS, SB | 2 | |
Mazandaran | unnamed brook | in Momey Khal | Ghaem Shahr | 760 | 36°04'24.0"N, 052°58'19.0"E | 11.5.2018 | JB, TS, SB | 1 | |
Mazandaran | Palang Darreh R. | SE of Shirgah | Shirgah | 320 | 36°16'31.0"N, 052°56'54.0"E | 11.5.2018 | JB, TS, SB | 2 | |
Mazandaran | LT of Palang Darreh R. | SE of Shirgah | Shirgah | 345 | 36°16'30.0"N, 052°56'51.0"E | 11.5.2018 | JB, TS, SB | 1 | |
Mazandaran | RT of Sardab Rud | SW of Kelardasht | Kelardasht | 2020 | 36°26'06.5"N, 051°03'52.6"E | 9.3.2018 | SB | 9 | |
Mazandaran | Sardab Rud | S of Vandarbon | Kelardasht | 2290 | 36°25'23.0"N, 051°02'12.4"E | 16.6.2018 | SB | 57 | |
Mazandaran | RT of Dohezar R. | N Holu Kaleh | Tonkabon | 880 | 36°37'37.5"N, 050°44'30.2"E | 16.6.2018 | SB | 83 | |
Mazandaran | Dohezar R. | SW Parde Sar | Tonkabon | 450 | 36°40'07.0"N, 050°49'20.0"E | 16.6.2018 | SB | 46 | |
Mazandaran | RT of Sehezar R. | S Parde Sar | Tonkabon | 570 | 36°38'41.5"N, 050°50'11.1"E | 16.6.2018 | SB | 20 | |
Mazandaran | Sehezar R. | S Parde Sar | Tonkabon | 540 | 36°39'01.0"N, 050°50'00.0"E | 24.8.2018 | SB | 21 | |
Mazandaran | Lavij Rud | SE of Kiakola | Noor | 820 | 36°21'33.1"N, 052°03'11.0"E | 24.8.2018 | SB | 8 | |
Mazandaran | Vaz Rud | E of Vaz Oliya | Noor | 1140 | 36°19'08.0"N, 052°08'24.1"E | 15.6.2018 | SB | 33 | |
Mazandaran | Safarud | SW of Ramsar | Ramsar | 490 | 36°52'55.8"N, 050°33'56.1"E | 15.6.2018 | SB | 1 | |
Mazandaran | LT of Safarud | SW of Ramsar | Ramsar | 610 | 36°53'24.4"N, 050°33'56.1"E | 15.6.2018 | SB | 2 | |
Mazandaran | LT of Safarud | SW of Ramsar | Ramsar | 330 | 36°54'06.1"N, 050°35'12.1"E | 15.6.2018 | SB | 11 | |
Mazandaran | Chalak Rud | SW of Galeshmahalleh | Ramsar | 100 | 36°49'13.0"N, 050°43'23.8"E | 15.6.2018 | SB | 3 | |
Mazandaran | LT of Chalak Rud | NW of Talesh Sara | Ramsar | 180 | 36°50'46.9"N, 050°40'25.9"E | 8.3.2018 | SB | 8 | |
Tehran | Ahar R. | near Igol | Fasham | 2020 | 35°55'11.2"N, 051°28'51.3"E | 8.5.2017 | AHS, MP, RJG, FN | 4 |
The most common Caucasiron species in the Alborz is E. (C.) cf. znojkoi distributed from the Talysh Mts. in the west to the Golestan NP in the east (Fig.
Three Caucasiron species, E. (C.) iranicus, E. (C.) alborzicus sp. nov., and E. (C.) shargi sp. nov., were described from the Alborz and are so far only known from there. E. (C.) iranicus is reliably reported from 12 localities, eight of them above 2000 m a.s.l. These include the Sabalan Mt. slopes in the western Alborz and the central Alborz, where it can co-occur with E. (C.) alborzicus sp. nov. (Fig.
Other streams investigated in Iranian mountain ranges were dominated by Baetidae, and Heptageniidae were generally only scattered there. Larvae of E. (C.) zagrosicus sp. nov. and E. (E.) zaitzevi were found only in five and seven localities respectively, relatively distant to each other in the Zagros (Fig.
We thank the Faculty of Natural Resources of the University of Gilan, the Department of Biodiversity and Ecosystem Management, Environmental Sciences Research Institute, Shahid Beheshti University, Tehran, and the Department of Environment, Natural History Museum and Genetic Resources, Tehran, for collaboration, support, sampling and export permits, and accompanying the authors during field trips. This research was part of the