A Taxonomic List of the Mayflies, Stoneflies and Caddisflies (Insecta: Ephemeroptera, Plecoptera and Trichoptera) of the Sikhote-Alin Biosphere Reserve



==Abstract==

Three orders of amphibiotic insects were examined from the eastern and western streams of the Sikhote-Alin State Nature Biosphere Reserve, which is located in the southern Russian Far East in the central portion of a mountain range with the same name. Data were obtained on Ephemeroptera, Plecoptera, Trichoptera (EPT) fauna inhabiting streams in the Sikhote-Alin State Nature Biosphere Reserve and its adjacent area. The data were collected by the author from 1980 to 2013. More than 30 thousand larvae and adult EPT were identified, and a systematic list of 220 species was formed. Among them, Ephemeroptera is represented by 63 species from 18 genera and 8 families. Plecoptera is represented by 61 species from 30 genera and 8 families. Trichoptera is represented by 96 species from 49 genera and 20 families. The EPT biodiversity study in the local protected areas is important for assessing the ecological preferences of aquatic organisms and understanding the formation of ecosystem structures under normal conditions, i.e., without anthropogenic influence. In addition, the EPT list is of great value, as it is widely used to control the quality of the environment via the EPT indicator index. The comprehensive list of Ephemeroptera, Plecoptera and Trichoptera (EPT complex) taxonomic species of Central Sikhote-Alin streams is submitted for the first time. In addition, a brief areal EPT fauna analysis is made.

==Keywords==

Amphibiotic insects ; EPT indicator index ; Sikhote-Alin State Nature Biosphere Reserve ; Southern Russian Far East ; Central Sikhote-Alin

==Introduction==

Sikhote-Alin State Nature Biosphere Reserve is located in the southern Russian Far East, in the central part of the Eponymous mountain range and on the shore of the Sea of Japan. Its territory is comprised of two clusters ([[#f0005|Fig. 1]] ), which encompass an area of 401.6 sq km. The mountainous terrain of the Sikhote-Alin mountain range is composed of numerous, complex overlapping mountain ridges. The average height of the mountain peaks is 800 m above sea level. The chaotic cluster of hills is dissected by river valleys, forming a dense (0.7 to 0.9 km per 1 sq km), extensive drainage system, with a variety of streams and basins. The rivers are less than 100 km in length, with average slopes of 50–100‰ and upper reach slopes of 150–200‰.

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[[Image:draft_García_174682007-1-s2.0-S2078152015000346-gr1.jpg|center|px|Terrain features of the study area.]]


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Fig. 1.

Terrain features of the study area.

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Rivers flowing from the eastern macroslope are fast, with frequent rapids and occasional small waterfalls. The water temperature in summer rarely rises above 16 °C. The river is more tranquil on the smoother, western macroslope, with water temperatures reaching 19–21 °C on hot days. The rivers are mainly fed by rainfall, which accounts for approximately 70–80% of the total annual river flow. According to various estimates, groundwater contributes to 5–20% of the flow, while snowmelt contributes to 10–20%.

The proximity to the ocean, location on the edge of the huge Eurasian continent and rugged terrain are the reasons for the unique climate of the region, which has a pronounced monsoon season. The annual cumulative temperatures of the growing season, which lasts approximately 150 days, amount to approx. 2000–2500 °С. The annual total precipitation on the eastern macroslope is 800 mm, and 650 mm on the western macroslope, with the majority (75–85%) occurring from April to November.

The watercourses of the Sikhote-Alin mountain range are of mountain and submontane types, with developed alluvial forms. These watercourses are of high commercial fishing value. During the freshwater portion of their lifecycle, juvenile salmon mainly feed on freshwater communities of aquatic insects, including mayflies (Ephemeroptera), stoneflies (Plecoptera) and caddisflies (Trichoptera) ([[#bb0035|Levanidov, 1969]] ). In addition, representatives of these orders are highly sensitive to various types of pollution and environmental changes. These characteristics are unique to these species, and make it possible for them to act as environmental indicators, including the widely used EPT biotic index ([[#bb0105|Semenchenko, 2004]] ). In this regard, EPT fauna identification is important for solving a number of fisheries issues, conducting water quality monitoring and assessing ecosystem states. In addition, the benthic communities sampled in this study do not suffer from anthropogenic influences, providing valuable data for the development of scientific methods for forecasting river ecosystem conditions based on various environmental disturbances.      

Despite the fact that mayflies, stoneflies and caddisflies in mountain and submontane streams and rivers are characterized by the highest species diversity, the insects in the streams of the reserve remained virtually unstudied until the 1980s. According to the literature, only 4 species of mayflies ([[#bb0005|Baikova, 1976]]  ;  [[#bb0110|Sinichenkova, 1981]] ), 4 species of stoneflies ([[#bb0150|Zhiltsova ''et al'' ., 1975                            ]] ; [[#bb0140|Zhiltsova and Levanidova, 1978]]  ;  [[#bb0100|Ricker, 1959]] ) and 16 taxa of caddisflies were known, 4 of which have not received any species affiliation ([[#bb0120|Timofeeva, 1978]] ).      

Based on a comprehensive study of the species compositions of these three orders of amphibiotic insects, the author has obtained lists of Ephemeroptera ([[#bb0050|Potikha, 1985]]  ;  [[#bb0070|Potikha, 2013]] ), Plecoptera ([[#bb0075|Potikha, 2014]] ; [[#bb0085|Potikha and Zhiltsova, 1996]] ; [[#bb0090|Potikha and Zhiltsova, 2001]]  ;  [[#bb0095|Potikha and Zhiltsova, 2005]] ) and Trichoptera ([[#bb0055|Potikha, 1991]] ; [[#bb0060|Potikha, 2001]] ; [[#bb0065|Potikha, 2002]]  ;  [[#bb0080|Potikha and Vshivkova, 2013]] ). The list is currently comprised of 220 taxa (Ephemeroptera — 63 taxa from 18 genera and 8 families; Plecoptera — 61 taxa from 30 genera and 8 families and Trichoptera — 96 taxa from 49 genera and 20 families). The complete taxonomic EPT list from watercourses of the Central Sikhote-Alin, as well as a brief areal analysis, is presented in the following sections.      

==Materials and Methods==

From 1980 to 2013, the author studied more than 90 streams and basins in the reserve and adjacent territories ([[#f0010|Fig. 2]] ). Over a thousand larvae samples and three thousand adult samples were collected and analyzed. In total, 30 thousand larvae and adults were identified, and over 2.5 thousand individual insects from scientific collections of academic institutions (ZI RAS, BSI FEB RAS) and MSU were studied. Imagoes were collected in coastal vegetation with traps, sweep nets (during swarming) and light traps. Larvae were collected from the surface of substrates via the forced drift method. Qualitative samples were preserved with 75% ethanol, and quantitative with 4% formalin. The samples are stored in the scientific collections of the Sikhote-Alin State Nature Biosphere Reserve, ZI RAS, BSI FEB RAS and MSU.

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[[Image:draft_García_174682007-1-s2.0-S2078152015000346-gr2.jpg|center|578px|Map of main sampling locations.]]


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Fig. 2.

Map of main sampling locations.

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Distribution types are listed according to [[#bb0145|Zhiltsova and Levanidova (1984)]] . Information from websites was used for compiling the taxonomic list ([[#bb0010|Ephemeroptera of The World]] ; [[#bb0040|Morse]]  ;  [[#bb0045|Order Plecoptera]] ). The taxonomic position of mayflies is listed according to [[#bb0025|Kluge, 2004]]  ;  [[#bb0030|Kluge, 2007]] . The Ephemerellidae family classification of [[#bb0020|Jacobus and McCafferty (2008)]]  was used. In addition, the Cinygmula classification of [[#bb0135|Wang and McCafferty (2004)]]  was utilized. The names of families, genera within families and species within genera are listed in alphabetical order.      

==Results and Discussion==

The taxonomic EPT complex list from the Sikhote-Alin State Nature Biosphere Reserve can be seen below, with statistics presented in [[#t0005|Table 1]] . The list includes 220 entries belonging to 97 genera and 36 families.

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Table 1.

The number of EPT complex taxa in the Sikhote-Alin State Nature Biosphere Reserve and adjacent territories.

|-

! Oder
! Families
! Genera
! Species
|-

| Ephemeroptera
| 8
| 18
| 63
|-

| Plecoptera
| 8
| 30
| 61
|-

| Trichoptera
| 20
| 49
| 96
|-

| Total
| 36
| 97
| 220
|}

==Taxonomic List==

Order '''Ephemeroptera'''

Family '''Ephemeridae'''

''Ephemera orientalis''  McLachlan, 1875      

''Ephemera strigata''  Eaton, 1892      

''Ephemera sachalinensis''  Matsumura, 1911      

Family '''Polymitarcyidae'''

''Ephoron shigae''  (Takahashi, 1924)      

Family '''Heptageniidae'''

''Cinygmula autumnalis''  Tiunova et Gorovaya, 2012      

''Cinygmula cava''  Ulmer, 1928      

''Cinygmula hirasana''  Imanishi, 1935      

''Cinygmula irina''  Tshernova et Belov, 1982      

''Cinygmula kurenzovi''  (Bajkova, 1965)      

''Cinygmula levanidovi''  Tshernova et Belov, 1982      

''Cinygmula putoranica''  Kluge, 1980      

''Cinygmula sapporensis''  (Matsumura, 1904)      

''Ecdyonurus''  (''Afghanurus'' ) ''aspersus''  Kluge, 1980      

''Ecdyonurus''  (''Afghanurus'' ) ''bajkovae''  Kluge, 1986      

''Ecdyonurus''  (''Afghanurus'' ) ''scalaris''  Kluge, 1983      

''Ecdyonurus''  (''Atopopus'' ) ''aurarius''  Kluge, 1983      

''Epeorus''  (''Proepeorus'' ) ''anatolii''  Sinitshenkova, 1981      

''Epeorus''  (''Belovius'' ) ''gornostajevi''  Tshernova, 1981      

''Epeorus''  (''Belovius'' ) ''ninae''  Kluge, 1995      

''Epeorus''  (''Belovius'' ) ''pellucidus''  (Brodsky, 1930)      

''Epeorus''  (''Belovius'' ) ''rubeus''  Tiunova, 1991      

''Epeorus (Iron) aesculus''  Imanishi, 1934      

''Epeorus (Iron) alexandri''  Kluge et Tiunova, 1989      

''Epeorus (Iron) maculatus''  (Tshernova, 1949)      

''Heptagenia''  (''Heptagenia'' ) ''sulphurea''  (Müller, 1776)      

''Rhithrogena bajkovae''  Sowa, 1973      

''Rhithrogena lepnevae''  Brodsky, 1930      

''Rhithrogena sibirica''  Brodsky, 1930      

Family '''Ameletidae'''

''Ameletus camtschaticus''  Ulmer, 1928      

''Ameletus cedrensis''  Sinitshenkova, 1977      

''Ameletus inopinatus labiatus''  Sinitshenkova, 1981      

''Ameletus longulus''  Sinitshenkova, 1981      

''Ameletus montanus arlecchino''  Kluge, 2007      

Family '''Siphlonuridae'''

''Siphlonurus immanis''  Kluge, 1985      

''Siphlonurus zhelochovtsevi''  Tshernova, 1952      

''Siphlonurus''  sp.      

Family '''Baetidae'''

''Baetis (Baetis) bicaudatus''  Dodds, 1923      

''Baetis (Baetis) fuscatus''  Linnaeus, 1761      

''Baetis (Baetis) pseudothermicus''  Kluge, 1983      

''Baetis (Baetis) ursinus ursinus''  Kazlauskas, 1963      

''Baetis (Baetis) vernus''  Curtis, 1834      

''Acentrella sibirica''  (Kazlauskas, 1963)      

''Baetiella tuberculata''  (Kazlauskas, 1963)      

''Сloeon''  sp.      

Family '''Leptophlebiidae'''

''Leptophlebia''  (''Neoleptophlebia'' ) ''japonica''  (Matsumura, 1931)      

''Leptophlebia''  (''Paraleptophlebia'' ) ''strandii''  Eaton, 1901      

''Leptophlebia''  sp. 1      

''Leptophlebia''  sp. 2      

Family '''Ephemerellidae'''

''Drunella aculea''  Allen 1971      

''Drunella cryptomeria''  (Imanishi 1937)      

''Drunella lepnevae''  Tshernova 1949      

''Drunella solida''  Bajkova 1980      

''Drunella triacantha''  Tshernova 1949      

''Ephemerella aurivillii''  Bengtsson 1908      

''Ephemerella''  f. ''thymalli''

''Ephemerella''  (''Hosoba'' ) ''atagosana''  Imanishi 1937      

''Ephemerella''  (''Zonadia'' ) ''kozhovi''  Bajkova 1967      

''Ephemerella''  (''Draeconia'' ) ''mucronata''  (Bengtsson 1909)      

''Cincticostella levanidovae''  (Tshernova 1952)      

''Cincticostella tshernovae''  (Bajkova 1962)      

''Serratella ignita''  (Poda 1761)      

''Serratella setigera''  (Bajkova 1967)      

''Serratella zapekinae''  Bajkova, 1967      

Order '''Plecoptera'''

Family '''Taeniopterygidae'''

''Taenionema japonicum''  (Okamoto, 1922)      

Family '''Nemouridae'''

''Zapada quadribranchiata''  (Zhiltzova, 1977)      

''Amphinemura borealis''  (Morton, 1894)      

''Amphinemura coreana''  Zwick, 1973      

''Amphinemura standfussi''  Ris, 1902      

''Amphinemura verrucosa''  Zwick, 1973      

''Nemoura arctica''  Esben-Petersen, 1910      

''Nemoura despinosa''  Zhiltzova, 1977      

''Nemoura geei''  Wu, 1929      

''Nemoura nigrodentata''  Zhiltzova, 1980      

''Nemoura papilla''  Okamoto, 1922      

''Protonemura ermolenkoi''  Zhiltzova, 1982      

Family '''Capniidae'''

''Capnia aligera''  Zapekina-Dulkeit, 1975      

''Capnia nearctica''  Banks, 1918      

''Capnia potikhae''  Zhiltzova, 1996      

''Capniella ghilarovi''  Zhiltzova, 1988      

''Capniella nodosa''  Klapálek, 1920      

''Eucapnopsis brevicauda''  (Claassen, 1924)      

''Isocapnia arcuata''  Zhiltzova, 1975      

''Isocapnia guentheri''  (Joost, 1970)      

''Isocapnia kudia''  Ricker, 1959      

''Paracapnia khorensis''  Zhiltzova, 1972      

''Paracapnia sikhotensis''  Zhiltzova, 1978      

''Paracapnia leisteri''  Zhiltzova et Potikha, 2005      

Family '''Leuctridae'''

''Leuctra fusca''  (Linnaeus, 1758)      

''Paraleuctra cercia''  (Okamoto, 1922)      

''Paraleuctra zapekinae''  Zhiltzova, 1974      

''Perlomyia levanidovae''  (Zhiltzova, 1975)      

''Perlomyia secunda''  (Zapekina-Dulkeit, 1955)      

''Perlomyia smithae''  Nelson et Hanson, 1973      

Family '''Pteronarcyidae'''

''Pteronarcys sachalina''  Klapálek, 1908      

Family '''Perlodidae'''

''Arcynopteryx dichroo''  (McLachlan, 1872)      

''Arcynopteryx polaris''  Klapálek, 1912      

''Diura majuscula''  (Klapálek, 1912)      

''Diura nanseni''  (Kempny,1900)      

''Megarcys magnilobus''  Zhiltzova, 1988      

''Megarcys ochracea''  Klapálek, 1912      

''Megarcys pseudochracea''  Zhiltzova, 1977      

''Pictetiella asiatica''  Zwick et Levanidova, 1971      

''Skwala compacta''  (McLachlan, 1872)      

''Stavsolus manchuricus''  Teslenko 1999      

''Isoperla eximia''  Zapekina-Dulkeit, 1975      

''Isoperla flavescens''  Zhiltzova et Potikha, 1986      

''Isoperla lunigera''  (Klapálek, 1923)      

''Isoperla maculata''  Zhiltzova, 1977      

Family '''Perlidae'''

''Kamimuria exilis''  (McLachlan, 1872)      

''Oyamia nigribasis''  Banks, 1920      

''Paragnetina flavotincta''  (McLachlan, 1872)      

Family '''Chloroperlidae'''

''Paraperla lepnevae''  Zhiltzova, 1970      

''Utaperla orientalis''  Nelson et Hanson, 1969      

''Alloperla deminuta''  Zapekina-Dulkeit, 1970      

''Alloperla mediata''  (Navás, 1925)      

''Alloperla rostellata''  (Klapálek, 1923)      

''Haploperla maritima''  Zhiltzova et Levanidova, 1978      

''Suwallia asiatica''  Zhiltzova et Levanidova, 1978      

''Suwallia decolorata''  Zhiltzova et Levanidova, 1978      

''Suwallia kerzhneri''  Zhiltzova et Zwick, 1971      

''Suwallia talalajensis''  Zhiltzova, 1976      

''Suwallia teleckojensis''  (Šámal, 1939)      

''Sweltsa illiesi''  Zhiltzova et Levanidova, 1978      

''Sweltsa lepnevae''  Zhiltzova, 1977      

Order '''Trichoptera'''

Family '''Apataniidae'''

''Apatania complexa''  (Martynov, 1935)      

''Apatania crymophila''  MacLachlan, 1880      

''Apatania zonella''  (Zetterstedt, 1840)      

Family '''Arctopsychidae'''

''Arctopsyche palpata''  Martynov, 1934      

Family '''Brachycentridae'''

''Brachycentrus americanus''  (Banks, 1899)      

''Brachycentrus japonicus''  Iwata, 1927      

''Micrasema''  (''gelidum'' ) ''primoricum''  Botosaneanu, 1990      

Family '''Ecnomidae'''

''Ecnomus tenellus''  (Rambur, 1842)      

Family '''Glossosomatidae'''

''Agapetus inaequispinosus''  Schmid, 1970      

''Agapetus levanidoruv''  Vshivkova et Morse, sp.n.      

''Electragapetus martynovi''  Vshivkova et Arefina, 1996      

''Electragapetus praeteritus''  Martynov, 1934      

''Glossosoma (Anagapetus) schmidi''  Levanidova, 1979      

''Glossosoma''  (''Synafophora'' ) ''altaicum''  Martynov, 1914      

''Glossosoma''  (''Synafophora'' ) ''angaricum''  (Levanidova, 1967)      

''Glossosoma''  (''Synafophora) intermedium''  (Klapálek, 1892)      

''Glossosoma''  (''Synafophora'' ) ''ussuricum''  (Martynov, 1934)      

Family '''Goeridae'''

''Archithremma ulachensis''  Martynov, 1935      

''Goera parvula''  Martynov, 1935      

''Goera squamifera''  Martynov, 1909      

''Goera tungusensis''  Martynov, 1909      

Family '''Hydrobiosidae'''

''Apsilochorema sutshanum''  Martynov, 1934      

Family '''Hydropsychidae'''

''Cheumatopsyche infascia''  Martynov, 1934      

''Hydropsyche orientalis''  Martynov, I934      

''Hydropsyche''  sp''.''

''Potamyia czekanovskii''  (Martynov, 1910)      

Family '''Hydroptilidae'''

''Hydroptila spinosa''  Arefina et Armitage, 2003      

''Orthotrichia tragetti''  Mosely, 1930      

''Oxyethira ecornuta''  Morton, 1893      

Family '''Lepidostomatidae'''

''Lepidostoma albardanum''  (Ulmer, 1906)      

''Lepidostoma elongatum''  Martynov, 1935      

''Lepidostoma sinuatum''  Martynov, 1935      

Family '''Leptoceridae'''

''Ceraclea excisa''  (Morton, 1904)      

''Ceraclea sibirica''  (Ulmer, 1906)      

''Mystacides bifida''  Martynov 1924      

''Mystacides sibirica''  Martynov 1935      

''Oecetis lacustris''  (Pictet, 1834)      

''Oecetis nigropunctata''  Ulmer, 1908      

''Triaenodes''  (''Ylodes'' ) ''levanidovae''  Morse et Vshivkova, 1997      

''Triaenodes unanimis''  MacLachlan, 1877      

Family '''Limnephilidae'''

''Asynarchus amurensis''  (Ulmer, 1905)      

''Brachypsyche rara''  (Martynov, 1914)      

''Brachypsyche sibirica''  (Martynov, 1924)      

''Brachypsyche''  sp. N      

''Chilostigma sieboldi''  MacLachlan, 1876      

''Chilostigmodes forcipatus''  Martynov, 1914      

''Dicosmoecus jozankeanus''  (Matsumura, 1931)      

''Ecclisomyia kamtshatica''  (Martynov, 1914)      

''Hydatophylax grammicus''  (MacLachlan, 1880)      

''Hydatophylax nigrovittatus''  (MacLachlan, 1872)      

''Hydatophylax soldatovi''  (Martynov, 1914)      

''Hydatophylax variabilis''  (Martynov, 1910)      

''Lenarchus productus''  (Morton, 1896)      

''Limnephilus alienus''  Martynov, 1914      

''Limnephilus correptus''  MacLachlan, 1880      

''Limnephilus fenestratus''  (Zetterstedt, 1840)      

''Limnephilus picturatus''  MacLachlan, 1875      

''Limnephilus quadratus''  Martynov, 1914      

''Limnephilus sericeus''  (Say, 1824)      

''Limnephilus stigma''  Curtis, 1834      

''Limnephilus''  sp. aff. ''subcentralis''  Brauer, 1857      

''Limnephilus tiunovae''  Arefina et Levanidova, 1996      

''Nemotaulius admorsus''  (MacLachlan, 1866)      

''Nemotaulius mutatus''  (MacLachlan, 1872)      

''Philarctus rhomboidalis''  Martynov, 1924      

''Pseudostenophylax amurensis''  (MacLachlan, 1880)      

Family '''Molannidae'''

''Molanna moesta''  Banks, 1906      

''Molannodes tinctus''  (Zetterstedt, 1840)      

Family '''Philopotamidae'''

''Dolophilodes''  (''Dolophilodes'' ) ''affinis''  Levanidova et Arefina, 1996      

''Dolophilodes mroczkowskii''  Botosaneanu, 1970      

''Kisaura aurascens''  (Martynov, 1934)      

''Wormaldia''  niiensis Kobayashi, 1985      

Family '''Phryganeidae'''

''Agrypnia czerskyi''  (Martynov, 1924)      

''Agrypnia picta''  Kolenati, 1848      

''Hagenella sibirica''  (Martynov, 1909)      

''Oligothricha lapponica''  (Hagen, 1864)      

''Semblis atrata''  (Gmelin, 1789)      

''Semblis phalaenoides''  (Linnaeus, 1758)      

Family '''Phryganopsychidae'''

''Phryganopsyche latipennis''  (Banks, 1906)      

Family '''Psychomyiidae'''

''Metalype uncatissima''  (Botosaneanu 1970)      

''Paduniella uralensis''  Martynov, 1914      

Family '''Rhyacophilidae'''

''Rhyacophila angulata''  Martynov, 1910      

''Rhyacophila coreana''  Tsuda, 1940      

''Rhyacophila depressa''  Martynov, 1910      

''Rhyacophila impar''  Martynov, 1914      

''Rhyacophila kardakoffi''  Navás, 1926      

''Rhyacophila kawamurae''  Tsuda, 1940      

''Rhyacophila lata''  Martynov, 1918      

''Rhyacophila lepnevae''  Levanidova, 1977      

''Rhyacophila monstrosa''  Levanidova et Schmid, 1977      

''Rhyacophila narvae''  Navás, 1928      

''Rhyacophila retracta''  Martynov, 1914      

''Rhyacophila sutchanica''  Schmid et Levanidova, 1986      

Family '''Stenopsychidae'''

''Stenopsyche marmorata''  Navás, 1920      

Family '''Uenoidae'''

''Neophylax relictus''  (Martynov, 1935)      

''Neophylax ussuriensis''  (Martynov, 1914)      

The EPT fauna of the Sikhote-Alin State Nature Biosphere Reserve are characterized by a large species diversity, which is representative of fauna from the Russian Far East. Ephemeroptera comprises approximately 36% ([[#bb0125|Tiunova, 2012]] ) of the fauna in the Russian Far East, while Plecoptera accounts for nearly 44% ([[#bb0115|Teslenko, 2007]] ) and Trichoptera roughly 24 ([[#bb0015|Ivanov, 2011]] ).      

The mayfly fauna in Sikhote-Alin State Nature Biosphere Reserve and adjacent territory watercourses are dominated by the Heptageniidae (24 species) and Ephemerellidae (15 species) families, followed by the Baetidae (8 species), Ameletidae (5 species) and Leptophlebiidae (4 species) families. The Ephemeridae and Siphlonuridae families are comprised of 3 species each, while Polymitarcyidae is represented by only one species.

The greatest number of stonefly species were in the Perlodidae (14 species), Chloroperlidae (13 species), Capniidae (12 species) and Nemouridae (11 species) families. The Leuctridae family includes 6 species, Perlidae 3 species and Taeniopterygidae and Pteronarcyidae only one species each.

The greatest number of caddisflies species were registered in the Limnephilidae (26 species) family, followed by Rhyacophilidae (12 species), Glossosomatidae (9 species), Leptoceridae (8 species) and Phryganeidae (6 species). Other families consist of 2–4 species, and the Arctopsychidae, Ecnomidae, Hydrobiosidae, Phryganopsychidae and Stenopsychidae families include only one species each.

Data on taxonomic composition should not be considered definitive, as ongoing research from Kolumbe River (Bolshaya Ussurka River basin), and in a new territory, suggests that the taxonomic list will be further updated with new species.

All amphibiotic insects identified up to the species level are summarized in two groups based on the type of distribution: the Holarctic and Palearctic groups ([[#t0010|Table '''2''']] ). In the Holarctic group, two types of habitats were identified, the circumpolar type, which widely covers the Palaearctic and Nearctic regions, and the amphi-Pacific type, which is spread throughout the East Asian Palearctic sector and north-western mountainous Nearctic regions, including a strip along the coast of the Arctic Ocean. In the second group, three types of habitats were identified, including the Trans Palearctic, East Palaearctic and Palearchaearctic groups. The Trans Palearctic group includes species that are both disjunctive and widespread in the Palearctic, while the East Palaearctic group combines species commonly found to the east of the Yenisei River and inhabiting areas along the west coast of the Pacific Ocean.

<span id='t0010'></span>

{| class="wikitable" style="min-width: 60%;margin-left: auto; margin-right: auto;"
|+

Table 2.

Biogeographical composition of the EPT fauna complex of the Sikhote-Alin State Nature Biosphere Reserve.

|-

! rowspan="2" | Distribution type
! colspan="2" | Mayflies
! colspan="2" | Stoneflies
! colspan="2" | Caddisflies
! colspan="2" | All groups
|-

! N
! %
! N
! %
! N
! %
! N
! %
|-

| colspan="9" | ''Holarctic group''
|-

| Circumpolar
| 1
| 1.7
| 3
| 4.9
| 13
| 14.0
| 17
| 8.0
|-

| Amphi-Pacific
| 1
| 1.7
| 2
| 3.3
| 1
| 1.1
| 4
| 2.0
|-

| colspan="9" | 
|-

| colspan="9" | ''Palaearctic group''
|-

| Palearchaearctic
| 
| 
| 
| 
| 
| 
| 
| 
|-

|  Mainland
| 10
| 17.2
| 20
| 32.8
| 14
| 15.0
| 44
| 20.7
|-

|  Mainland-island
| 7
| 12.1
| 5
| 8.2
| 17
| 18.3
| 29
| 13.7
|-

|  East Palaearctic
| 33
| 56.9
| 28
| 45.9
| 30
| 32.3
| 91
| 42.9
|-

|  Trans Palearctic
| 6
| 10.4
| 3
| 4.9
| 18
| 19.3
| 28
| 12.7
|-

| Total
| 58
| 
| 61
| 
| 93
| 
| 212
| 
|}

Remark. N — number of taxa identified up to the species level.

In general, the EPT species of rivers and water basins in the Sikhote-Alin State Nature Biosphere Reserve are heterogeneous in biogeographic composition ([[#t0010|Table '''2''']] ). They are characterized by a high number of Palaearctic group species, nearly 90.0% (191 species). Within the Palaearctic group, the contribution of East Palaearctic species is the most significant at 42.9% (91 species). These are followed by species within the Palearchaearctic distribution (34.4%; 73 species), of which a portion are characterized as mainland subregion distribution dominate (20.7%; 44 species). Species within the Trans Palearctic distribution type occupy a subordinate position at 12.7% (28 species). The Holarctic group is not numerous and is represented by a total of 21 species (10.0%).      

A diversity can be observed in the species correlations within the East Palaearctic and Palearchaearctic habitat types. For example, among mayflies, East Palaearctic species significantly dominate over Palearchaearctic species, constituting 56.9% and 29.3%, respectively. Among stoneflies and caddisflies, the species correlations within the East Palaearctic and Palearchaearctic habitat types are insignificant. However, stoneflies are dominated by East Palaearctic species (45.9%), while caddisflies are dominated by Palearchaearctic species (33.3%). Stoneflies (20 species) and mayflies (10 species) prevail among Palearchaearctic species, the distribution of which is limited to the mainland subregion. Species common in the mainland-island subregion are predominantly caddisflies (17 species). Palearctic species, which are widely distributed, constitute only 4.9% of all stoneflies, while their portions of mayflies and caddisflies are higher, at 10.4% and 19.3%, respectively. Only two mayfly species are registered in the Holarctic group (3.4%), including circumboreal ''Ephemera mucronata''  and amphi-Pacific ''Baetis''  (B.) ''bicaudatus'' . Stoneflies are represented by five species (8.2%), and among them there are three species with amphi-Bering distribution and two species with amphi-Pacific distribution. Caddisfly species are the most abundant in the Holarctic group (14 species, 5.1%).      

In addition, three species, ''Ecnomus tenellus'' , ''Molanna moesta''  and ''Oecetis lacustris'' , reach beyond the Holarctic region in their distribution. The first species is found in the Oriental and Afrotropical regions, and the other two in the Oriental region ( [[#bb0040|Morse]] ). This species correlation by habitat type is typical of the southern Russian Far East and reflects the general fauna formation laws in this region ( [[#bb0130|Vshivkova, 1995]] ; [[#bb0110|Sinichenkova, 1981]]  ;  [[#bb0125|Tiunova, 2012]] ).      

In summary, this study of taxonomic EPT complex diversity at a local scale not only provides valuable information for solving biogeographic problems but also serves as a basis for assessing the ecological preferences of aquatic organisms and understanding the formations of ecosystem structures in a natural environment. Although the Reserve is free of human influence, the results can also be used for monitoring areas subjected to anthropogenic stress.

==Acknowledgments==

The author would like to thank the staff of the Sikhote-Alin State Nature Biosphere Reserve — M.V. Podushko, G.P. Averkova, S.V. Elsukov, N.V. Kholushkina and N.V. Vasilenko for the interesting findings. Special thanks to the staff of BSI FEB RAS — T.M. Tiunova, L.A. Teslenko, T.S. Vshivkova and T.I. Arefina-Armitage, as well as ZI RAS employee L.A. Zhiltsova, for their invaluable advice and assistance in identifying the samples.

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