Research Article |
Corresponding author: Bazartseren Boldgiv ( boldgiv@num.edu.mn ) Academic editor: Mariano Michat
© 2019 Davaadorj Enkhnasan, Bazartseren Boldgiv.
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:
Enkhnasan D, Boldgiv B (2019) Biogeography of predaceous diving beetles (Coleoptera, Dytiscidae) of Mongolia. ZooKeys 853: 87-108. https://doi.org/10.3897/zookeys.853.33908
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The bio-geographical composition and spatial distribution patterns of dytiscid assemblages in Mongolia are relatively unexplored. In this study, we compiled a list of 99 dytiscid species belonging to 20 genera and five subfamilies recorded in Mongolia and investigated species richness, spatial distribution and bio-geographical composition of the Mongolian dytiscid fauna. This study encompasses the information of currently recorded species and their geographic localities in Mongolia based on our own data and literature sources. We examined how dytiscid species richness was related to sub-basins of surface water network, as well as to geographical elevations within Mongolia. The majority of the Mongolian dytiscid fauna was associated with the sub-basins belonging to Arctic Ocean (80 species, 80.8%) and Central Asian Inland (60 species, 60.6%) basins. Only a few species of dytiscids belonged to the remaining river basins. Species richness of dytiscids and total area of sub-basins were not correlated, but species composition of dytiscids differed significantly among the sub-basins.
We observed that most of the species (77 species or 77.8% of total fauna) were recorded in a wide range of elevations and mid-altitudes (1000–2000 m a.s.l.) and showed the greatest diversity of dytiscids. Regarding the bio-geographical composition, species with wide geographical distributions (27.3% of dytiscids), were Palearctic species, while species of Arctic origin (21.2%) together with Boreal elements (16.2%) comprised a large proportion of the dytiscid fauna in Mongolia.
Geographical distribution, altitudinal pattern, dytiscid, range, river sub-basin
Under global change, natural ecosystems in Mongolia are experiencing greater-than-global average rate of climate change, as well as shifting anthropogenic influences. The country has a large landlocked territory covering an area of 1,564,118 square kilometers in Inner Asia. It is located on the Mongolian Plateau with an average elevation of 1580 m (range of elevation: 560–4374 m), and about 85% of its area lies over 1000 m above sea level (
The country has a large variety of geographic features including high mountains in the west, forests in the north, deserts in the south and plain steppes in the central and eastern regions, with various environmental and geographic formations. Insect biogeographic studies have been done only for a few groups of terrestrial insects of Mongolia (
A large number of water beetles were collected in June–July in 2003–2006 and 2008–2011 during the Selenge River Basin and Mongolian Aquatic Insect Survey Expeditions, respectively, as well as in 2009–2015 by Mongolian and Russian researchers within the framework of the Mongolian-Russian Biological Expedition.
Several works on dytiscids have been published based on these studies (
Although all of the above-mentioned studies pointed out only general distributions and taxonomy of the dytiscids; a comprehensive overview of the zoogeography of the Mongolian aquatic Coleoptera has not yet been done. In this paper, we attempt to make bio-geographical analyses of dytiscids known for Mongolia, in relation to the country’s surface water network and geographical features.
Mongolia has an extreme continental climate, with four distinctive seasons. Temperature fluctuates greatly, both daily and annually, with low rainfall (
Our database consists of dytiscid samplings from ten sub-basins belonging to three water basins. Mongolia is situated on three international river basins (
Watersheds and sub-basins of Mongolia. Gray dots represent all sampling points of dytiscids. Abbreviations: KZ – Kazakhstan; WS – West Siberia; ES – East Siberia; FE – Far East; MAN – Manchuria; NMO – Nei Mongol; GAN – Gansu; XIN – Xinjiang; AOB–Arctic Ocean Basin; POB – Pacific Ocean Basin; CAIB – Central Asian Inland Basin.
Rivers belonging to AOB are most extensively developed in the north and constitute the country’s major river system, i.e., the Selenge River system, which drains via Lake Baikal to the Arctic Ocean. Some tributaries of Siberia’s Yenisei River, which also flows to the Arctic Ocean, originate in the mountains of northwestern Mongolia (
Many rivers of western Mongolia end at lakes in the CAIB, most often in those of the Great Lakes Depression. The depression is named so because it contains six major Mongolian lakes: the saline Uvs Nuur, Khyargas Nuur, Durgun Nuur and the fresh water Khar-us Nuur, Khar Nuur and Airag Nuur, as well as number smaller ones. The major rivers are Khovd, Zavkhan, and Tes. The few streams of southern Mongolia also do not reach the sea but run into lakes or deserts (
In northeastern Mongolia, rivers of POB such as Onon, Kherlen and Khalkh gol River drain into the Pacific after joining the Shilka River in Russia and the Amur (Heilong Jiang) rivers, forming the tenth longest river system in the world (
There are about 4113 rivers in Mongolia, with a total length of 67,000 km. The longest river is the Orkhon at 1124 kilometers in length. Large rivers originate in the mountainous areas in the north and west of the country – primarily in the Mongol Altai, Khangai-Khuvsgul and Khentii mountain ranges – where small rivers and mountain streams merge to create well-developed water networks. There are also over 3000 big and small lakes, 6900 springs, 190 glaciers and 250 mineral water springs in the country (
The surface water network is of greatest density in the north of the country. In contrast, the southern, central and southeastern parts of the country have few rivers or other surface water resources. In the interior drainage basins, in the western and southern areas of Mongolia, seasonal or intermittent streams end in salt lakes or disappear into the desert. The rivers’ main water sources are rainfall, groundwater, snow and glaciers, with melting snow accounting for 15–20 percent of the annual runoff (
During our study, we collected 3517 beetle specimens from 630 sampling points (Fig.
The relative area of the water sub-basin of Mongolia was taken from the classification of Mongolian water resources (
Predaceous diving beetle collections are currently deposited in the Laboratory of Entomology, Institute of General and Experimental Biology, Ulaanbaatar, Mongolia.
We have classified the dytiscid species of Mongolia in accordance with the earlier systematic work of
Palearctic. Species distributed throughout the whole Palearctic Region.
Holarctic. Inhabitants of the northern regions of the European Russia and Siberia (to Kamchatka i.e., tundra and taiga).
Oriental. Occurring at the border between Palearctic and Oriental regions: India and Pakistan, Kashmir, Himachal Pradesh, Uttar Pradesh, Nepal, Sikkim, and Darjeeling, Bhutan, Arunachal Pradesh.
Arctic. Species of Arctic origin with occurrence in the tundra and the northern edge of the taiga. They are distributed in the south to Transbaikalia and in Western Europe to northern Sweden and Norway.
Boreal. This group is the largest. They live in the taiga and insular forests of Eurasia, in the plains. They occur in the northern and central belt of European Russia, in Siberia and highlands of the Caucasus; they also occur in northern and central Europe to eastern France and northern Italy.
Mediterranean. Species widely distributed throughout the Mediterranean (southern Europe, North Africa, and Asia Minor); east Mediterranean species occur in the Balkans, Asia Minor, Syria and western Iran.
Steppe. Species of the steppe zone of European Russia, western Siberia, northern Kazakhstan, eastern Transcaucasia, Turkmenia (some species of this group reach Hungary and Austria in the west).
Turanian. Species occurring in the mountains of central Asia, Sinkiang, Tien Shan.
Palearchearctic. Species from Korea, China and Japan.
Similarities of dytiscid assemblages among the sub-basins were calculated using the Bray-Curtis’ quantitative formula (Bray and Curtis 1957) and the Simple Average Linkage for hierarchical clustering of objects. The results obtained were presented in a similarity dendrogram. Similarities among objects were determined using Biodiversity Pro v.2 software (McAleece et al. 1997). Square-root transformation was used to meet the assumption of normality because the data were counts (
Based on our investigations, the dytiscid fauna of Mongolia comprises 99 species belonging to 20 genera in five subfamilies. A list of dytiscid species and their occurrences in various sub-basins of Mongolia are given in Appendix
The greatest diversity of dytiscids was recorded from the Selenge River Basin and Depression of Great Lakes. The majority of dytiscid species of Mongolia was represented by two subfamilies, Hydroporinae (44 species) and Agabinae (36), that altogether comprise about 80 percent of the total dytiscid species. The other three subfamilies were Dytiscinae (11), Colymbetinae (7) and Laccophilinae (1), which were only represented by a few species in Mongolia (Appendix
The most species-rich and commonly encountered genera in Mongolia were Agabus (25 species), Hygrotus (14), Hydroporus (14), Ilybius (10), Nebrioporus (4), Graphoderus (4) and Rhantus (4). The other genera included less than four species each. Colymbetes, Dytiscus, Hydroglyphus and Oreodytes were each represented by three species, while Hydaticus, Acilius and Bidessus had two species each. Six genera were represented by a single species, namely Boreonectes, Laccophilus, Laccornis, Nectoporus, Platambus and Zaitsevhydrus. Thus, few genera were species-rich, whereas the majority comprised fewer species, with the mean number per genus = 5.9 species.
Because China and Russia are large countries and have many diverse zones geographically, our analysis also focused on species composition of surrounding regions in these countries adjacent to Mongolia, in order to reveal species which are shared among them. Fauna of dytiscids in the closest seven regions of Russia and China, as well as Kazakhstan were included. Based on information of the distribution of 261 species, 27 genera of dytiscids were compiled from adjacent regions and Mongolia and a presence or absence matrix for species in these regions was constructed. These analyses found that faunistic similarity coefficients between Mongolia and adjacent regions ranged from 7.1% to 58.4% (Fig.
Generic and species richness of dytiscids in Mongolia and its surrounding regions. Abbreviations: GAN – Gansu; MAN – Manchuria; NMO – Nei Mongol; XIN – Xinjiang; MNG – Mongolia; FE-Russian Far East; ES – East Siberia; WS – West Siberia; KZ – Kazakhstan. Source: Catalogue of Palearctic Coleoptera (
Similarity index of the Mongolian dytiscid fauna with neighboring regions.
GAN | MAN | NMO | XIN | FE | ES | WS | KZ | |
---|---|---|---|---|---|---|---|---|
MNG | 13.0 | 7.1 | 18.9 | 15.9 | 45.3 | 58.4 | 49.0 | 39.8 |
GAN | * | 0.0 | 25.9 | 24.6 | 10.1 | 10.9 | 4.4 | 10.6 |
MAN | * | * | 4.7 | 7.4 | 9.1 | 6.8 | 9.7 | 3.9 |
NMO | * | * | * | 20.3 | 25.2 | 18.0 | 11.5 | 10.3 |
XIN | * | * | * | * | 10.5 | 8.3 | 9.3 | 28.1 |
FE | * | * | * | * | * | 48.3 | 33.7 | 22.2 |
ES | * | * | * | * | * | * | 67.3 | 45.8 |
WS | * | * | * | * | * | * | * | 54.5 |
Most of the species recorded for the dytiscid fauna of Mongolia (86 species) were common with those of the surrounding regions. The East (similarity index 58.4%, 60 species shared) and West Siberia (51 species shared) regions of Russia showed the closest similarity with the Mongolian dytiscid fauna. The Manchuria (7.1%, four species shared) region of China had the lowest faunal similarity with Mongolia (Fig.
Dytiscids in the 10 sub-basins occurred with various numbers of species, from 5 to 79. Each of these sub-basins had a peculiar composition of dytiscids, but there were several dominant species in most of the sub-basins. Seventy-nine species were found from Selenge River Basin, 45 species from the Depression of Great Lakes, 26 species from Gobi Basin, 22 species from Tes River Basin and 21 species from Shishkhed and Valley of Lakes River Basins. The species richness of dytiscids in other sub-basins varied between 5 and 17 (see Appendix
Only Hygrotus marklini Gyllenhal, 1813 was common to nine sub-basins, six other species, namely H. impressopunctatus Schaller, 1783, H. flaviventris Motschulsky, 1860 Agabus adpressus Aubé, 1837, Laccophilus biguttatus, Oreodytes mongolicus Brinck, 1943 and Rhantus frontalis Marsham, 1802 inhabited seven out of 10 sub-basins, which provide a variety of habitats. While other five species, A. coxalis Sharp, 1880, R. notaticollis Aubé, 1837, Dytiscus dauricus Gebler, 1832, Nebrioporus airumlus Kolenati, 1845 and Hydroporus acutangulus complex Thomson, 1856 were found in six sub-basins. Agabus dichrous Sharp, 1878, Hygrotus unguicularis Crotch, 1874, Ilybius poppiusi Zaitzev, 1907, Nectoporus sanmarkii Sahlberg, 1826 and O. septentrionalis Gyllenhal in C.R. Sahlberg, 1826 were recorded from five sub-basins, while nine other species, A. infuscatus Aubé, 1838, A. pallens Poppius, 1905, Graphoderus austriacus Sturm, 1834, H. enneagrammus Ahrens, 1833, H. nigrolineatus Steven in Schönherr, 1808, I. cinctus Sharp, 1878 and I. lateralis Gebler, 1832 were distributed in four sub-basins. The other 32 species were found in two or three sub-basins. From our data it is apparent that 43 species have restricted ranges within Mongolia, because of their distribution being restricted to a single sub-basin (see Appendix
Based on the information of the distribution of 99 dytiscid species from different river basins in Mongolia, a presence or absence matrix for species in the nine regions was constructed, except BRB because there were only five species recorded so far (See Appendix
A dendrogram depicting the dytiscid faunal similarity among the water sub-basins of Mongolia. DGLB – Depression of Great Lakes Basin; GRB – Gobi River Basin; TRB – Tes River Basin; VLRB – Valley of Lakes River Basin; KhGRB – Khalkh Gol River Basin; KhRB – Kherlen River Basin; ORB – Orkhon River Basin; SRB – Selenge River Basin; ShRB – Shishkhed River Basin; BRB – Bulgan River Basin.
It should be noted that because of different sampling effort, the diversity of dytiscids reported here for some basins, such as the Valley of Lakes, Bulgan River Basin and Gobi Basin etc. might not be fully representative of reality. Overall, the dytiscid fauna of the various basins was relatively distinct, confirming the well-established classification of the sub-basins in Mongolia.
Numbers of dytiscid species of different zoogeographical origins listed for 10 sub-basins of Mongolia.
Bio-geographical ranges | AOB | CAIB | POB | |||||||
---|---|---|---|---|---|---|---|---|---|---|
SRB | ShRB | BRB | DGLB | GRB | TRB | VLRB | KhGRB | KhRB | ORB | |
Arctic origin | 19 | 7 | 1 | 13 | 5 | 4 | 6 | 2 | 5 | 4 |
Boreal | 13 | 5 | 2 | 9 | 3 | 5 | 4 | 2 | 3 | 5 |
Holarctic | 1 | 1 | 0 | 1 | 0 | 1 | 2 | 0 | 0 | 0 |
Turanian | 5 | 3 | 0 | 4 | 3 | 2 | 2 | 0 | 1 | 1 |
Mediterranean | 0 | 0 | 0 | 0 | 2 | 1 | 0 | 0 | 0 | 0 |
Oriental | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
Palearchearctic | 2 | 0 | 0 | 2 | 1 | 0 | 1 | 0 | 1 | 1 |
Palearctic | 32 | 4 | 1 | 10 | 7 | 7 | 5 | 0 | 4 | 2 |
Steppe zone | 7 | 1 | 1 | 6 | 5 | 4 | 1 | 5 | 3 | 2 |
Overall species | 79 | 21 | 5 | 45 | 26 | 23 | 21 | 9 | 17 | 16 |
Given the geographic distribution in various sub-basins of Mongolia reported above, there were some mismatches between the area of each basin and the respective number of dytiscid species. The highest number of species (79 of a total of 99 species) was recorded in the Selenge River Basin, though the total area of this region is only 18.9% of the territory of Mongolia. Forty-five species were found from the Depression of Great Lakes that covers 16.0% of the country. Twenty-one and twenty-six species were recorded in Shishkhed and Gobi basins, which comprise 1.3% and 39.9 % of the entire area of the country, respectively. Species richness of dytiscids varied also among the sub-basins. One-way ANOVA showed that significant differences were observed in the species richness (F9, 1077 = 4.34; p<0.0001) among the sub-basins. However, there was no significant relationship between the species richness of dytiscids and the total area of each sub-basin (r = 0.46, p = 0.1708).
Our analysis of patterns of dytiscid species distribution with altitudes was based upon collection data from 1077 different elevation points. Dytiscid community composition was different by altitudes, as most of the species were recorded in a wide range of elevations (Fig.
The number of species observed in the various altitude ranges differed significantly (p = 0.0006). Twelve species, namely Acilius sulcatus Nicolai, 1822, Agabus kaszabi Guéorguiev, 1972, A. moestus Curtis, 1835, Bidessus nasutus Sharp, 1887, Colymbetes pseudostriatus Nilsson, 2002, Hydaticus aruspex Clark, 1864, Hygrotus inaequalis Fabricius, 1777, Hydroporus palustris Linnaeus, 1760, Ilybius cinctus Sharp, 1878, Ilybius erichsoni Gemminger & Harold, 1868, Laccornis oblongus Stephens, 1835, Platambus maculatus Linnaeus, 1753 occurred in the range of 801–1000 m a.s.l., while Graphoderus zonatus verrucifer Sahlberg, 1824, Hydroporus angusi Nilsson, 1990, H. fuscipennis Schaum & Kiesenwetter, 1867, Ilybius balkei Fery & Nilsson, 1993 and I. opacus Aubé, 1837 were restricted to lower altitudes, between 600 and 800 m a.s.l..
Several other species (i.e. Agabus costulatus Motschulsky, 1859, A. lineatus Gebler, 1848, Boreonectes aff. emmerichi Falkenström, 1936, Hydroporus morio Aubé, 1838 and H. notabilis LeConte, 1850) were reported at elevations of 2001 – 2300 m a.s.l..
Species richness was greatest between 1000 and 2000 m a.s.l., as 75 species (77.8% of total diversity) was recorded in this mid-altitude range. Thus, we found a peak of species richness between 1400 and 1600 m a.s.l.
The geographic distribution of all known species of dytiscids in Mongolia was compiled, and the species were divided into groups based upon their range. Species of Boreal (16 species, 16.8%) and Arctic origin (21 species, 22.1%) comprised a large proportion of the dytiscid fauna, due to the extremely harsh and fluctuating climate of Mongolia (Fig.
Relationship of generic and species diversity of dytiscids in the water sub-basins of Mongolia according to their biogeographic regions. Abbreviations: AO – species of Arctic origin; BO – Boreal species; HO – Holarctic species; TU – Turanian species; ME – Mediterranean elements; OR – Oriental; PLA – Palearchearctic; PL – species distributed throughout the whole Palearctic; SZ – species of the Steppe zone.
Two species, Oreodytes mongolicus and Agabus kaszabi were found to be endemic to Mongolia. Although O. mongolicus was common throughout the whole country, it was collected mainly from sub-basins DGLB, TRB and SRB. Twenty-seven were Palearctic species (27.3% of total), namely Acilius sulcatus, Agabus blatta, A. congener, Agabus kaszabi, A. kholini, A. laferi, Agabus svenhedini, A. udege, Bidessus unistriatus, Boreonectes aff. emmerichi, Colymbetes pseudostriatus, Graphoderus cinereus, Hydaticus continentalis, Hydroglyphus licenti, Hydroporus angusi, H. crinitisternus, H. kabakovi, H. palustris, H. uenoi, Hygrotus chinensis, H. inaequalis, H. parallellogrammus, Ilybius balkei, I. chishimanus, Oreodytes shorti, Platambus maculatus and Rhantus vermiculatus. Twenty-one were Arctic species (22.1%), namely Agabus adpressus, A. aequalis, A. arcticus alpinus, A. costulatus, A. coxalis, A. lapponicus, A. moestus, A. thomsoni, Colymbetes dahuricus, C. dolabratus, Dytiscus latro, Hydroglyphus hamulatus, Hydroporus acutangulus complex, H. sibiricus, H. submuticus, Hygrotus unguicularis, Ilybius lateralis, I. obtusus, I. poppiusi and Oreodytes mongolicus. Sixteen Boreal species (16.1%), such as Acilius canaliculatus, Agabus biguttulus, A. discolor, Graphoderus zonatus verrucifer, G. zonatus zonatus, Hydroglyphus geminus, Hydroporus elongatulus, Hygrotus marklini, H. quinquelineatus, Ilybius angustior, Laccornis oblongus, Nebrioporus assimilis, N. depressus, Oreodytes septentrionalis, Rhantus frontalis, and R. notaticollis were also found to be widely distributed in Mongolia.
The quantitatively most important bio-geographical elements were Holarctic (14 species, 14.1%), Steppe (11 species, 11.1%) and Turanian (5 species, 5.0%) species. Mediterranean (Hygrotus confluens and Agabus nebulosus) and Palearctic species (Dytiscus dauricus, Nebrioporus hostiles) were each represented by two species. Finally, only one species from the Oriental Region (Agabus japonicus continentalis) was recorded from Mongolia.
Mongolia has representatives of about 1.12% of the known world dytiscid genera and 2.3% of the currently described species (
The Agabinae and Hydroporinae are the largest dytiscid subfamilies in the world and the dominant groups in most habitats. The prevalence of these subfamilies has been reported to increase with increasing variety of water bodies (
More than half of the recorded species in the Mongolian dytiscid fauna belong to the genera Agabus, Hygrotus, Hydroporus and Ilybius. Other genera containing three or more species in Mongolia were: Dytiscus (3 species), Graphoderus (4), Nebrioporus (4), Oreodytes (3), Rhantus (4), Colymbetes (3) and Hydroglyphus (3). Together, the eleven most diverse genera constituted nearly 88.5% of the dytiscid species known from Mongolia, while other genera, such as Acilius, Hydaticus, Bidessus, Boreonectes, Laccophilus, Laccornis, Nectoporus, Platambus and Zaitsevhydrus comprised a much smaller proportion of the fauna. Similar faunistic patterns were found in other regions surrounding Mongolia, e.g., Russian Far East and northern China (
The “Arctic Ocean Basin” group encompasses the Selenge, Shishkhed and Bulgan River Basins. The “Pacific Ocean Basin” group includes the Kherlen, Onon and Khalkh Gol River Basin. The “Central Asian Inland Basin” group consists of the Tes, Depression of Great Lakes, Valley of Lakes and Gobi Basin. The results show that the faunal composition of dytiscids is more similar among the sub-basins due to geographical adjacency to one another, and reflect the main landscape pattern of Mongolia. In particular, the similarity of dytiscid fauna between AOB and CAIB was 36.5% and between CAIB and POB 28.6%.
It is notable that the most common species (e.g. Hygrotus impressopunctatus, H. marklini, Agabus adpressus, A. coxalis, Oreodytes mongolicus, O. septentrionalis, Rhantus notaticollis, Dytiscus dauricus, Hydroporus acutangulus complex, Hygrotus flaviventris, Laccophilus biguttatus) tended to be widely distributed across various sub-basins, but in contrast the uncommon and rare species (Agabus biguttulus, A. clavicornis, A. congener, A. angusi, A. kholini, A. laferi, A. lapponicus, A. lineatus, Colymbetes dahuricus, C. pseudostriatus, Graphoderus cinereus, G. zonatus verrucifer, Hydroglyphus licenti, Hydroporus angusi, H. elongatulus, H. fuscipennis, H. kabakovi, H. morio, H. nigellus, H. notabilis, H. palustris, H. submuticus, H. uenoi, Hygrotus flaviventris, H. urgensis, I. balkei, I. chishimanus, I. erichsoni, I. opacus and Platambus maculatus) were highly restricted in their distribution, generally occurring only in one basin.
Aquatic macro-invertebrate assemblages can be affected by various local and regional environmental factors, such as chemical and physical characteristics of stream water, hydrology, and geographic location, as well as climatic factors.
Generally, the species richness of local dytiscid communities is primarily influenced by climatic conditions (e.g. temperature regimes, precipitation), landform, and microhabitat patterns (e.g. vegetation cover, erosion, variety of water bodies). In Mongolia, however, due to the high habitat heterogeneity, species diversity of dytiscids in various sub-basins may differ as a consequence of water physico-chemical parameters that can determine whether a species is present or absent within a locality; thus, small-scale patterns of habitat distribution are important for dytiscids in Mongolia, especially in arid regions.
We suggest that the clear differentiation of elevational distribution observed for the majority of dytiscid species considered in this study is due to local geographic relief, as the majority of the country exhibits mountainous landscapes (about 85% of its area is over 1000 m a.s.l.), the exception being the plain grasslands of eastern Mongolia.
The mid-elevation peak in dytiscid diversity is sometimes attributed to the warmer and better wetland habitat conditions and prey resource availability at these elevations. Based on research from various regions, it has been observed that high diversity of diving beetles depends on the number of wetland types represented in a landscape, and thus it is possible to achieve high diversity in a small area by combining permanent and temporary wetlands, as well as systems located in wooded and open environments (
The biogeographic composition of dytiscid fauna in Mongolia confirms that it is one of the representative parts of the Palearctic dytiscid fauna. The majority of dytiscid species in Mongolia are widespread in the whole Palearctic region, with the addition of Holarctic elements. Thus, species of Arctic and Boreal regions are widely distributed in Mongolia and comprise more than one third of the total number of species (37.4%). The other specific characteristic of the Mongolian dytiscid fauna is the presence of species from the Oriental Region, as for instance Agabus japonicus continentalis.
Finally, it should be noted that the magnitude of climatic changes in temperature and precipitation are predicted to stress a variety of ecosystems directly or indirectly. Most attention has focused on how climate change will affect terrestrial ecosystems, but aquatic ecosystems (e.g., ponds, lakes, streams, and rivers) will also experience parallel changes in diel, seasonal, and annual temperature and precipitation patterns. Therefore, we need to focus on issues related to the effect of increased temperature on the characteristics of biogeographical distribution of dytiscids. Detailed biogeographical surveys play an important role in providing information of what species are present in sub-basins and understanding their ecological roles, to better manage and protect aquatic ecosystems for the future.
We would like to thank Dr. Jon Gelhaus (project leader), Dr. Balgan Namkhaidorj and all members of “Selenge River Basin” and “Mongolian Aquatic Insect Survey (MAIS)” project for their help in collecting dytiscid specimens and sorting samples of water beetles. This research was supported by the US National Science Foundation grants (‘‘Survey and Inventory of the Aquatic Macro invertebrates of the Selenge River Drainage Basin, Mongolia’’ DEB-0206674; ‘‘Survey and Inventory of the Aquatic Insects of the Altai and Hangai Mountains’ Drainages, Mongolia’’ DEB-0743732). BB was supported by the Taylor Family-Asia Foundation Endowed Chair in Ecology and Conservation Biology. We thank Drs. Manfred Jäch and Helen Shaverdo, Natural History Museum of Vienna, Austria, for granting us the opportunity to work on dytiscid collections at that museum, and helping with identification and verification of some dytiscids, respectively. We would like to thank the reviewer(s) for careful and thorough reading, thoughtful comments and constructive suggestions, which helped to improve the quality of this manuscript.
Compiled dytiscid species list from own data and literature sources, and their distribution in sub-basins of Mongolia.
ID | Species name | DGLB | GRB | TRB | VLRB | KhGRB | KhRB | ORB | SRB | ShRB | BRB | Total occurrence in the basins | Total abundance of species |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|
Agabinae | |||||||||||||
1 | Agabus aequalis Sharp, 1882 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 3 |
2 | *Agabus amoenus Solsky, 1874 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 5 |
3 | *Agabus angusi Harris, 1828. | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 1 | 1 |
4 | Agabus arcticus alpinus Motschulsky, 1860 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 1 | 0 | 3 | 21 |
5 | Agabus congener Thunberg, 1794 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 1 |
6 | Agabus costulatus Motschulsky, 1859 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 2 | 24 |
7 | Agabus coxalis Sharp, 1880 | 1 | 1 | 0 | 0 | 1 | 0 | 1 | 1 | 1 | 0 | 6 | 21 |
8 | Agabus infuscatus Aubé, 1838 | 1 | 0 | 0 | 1 | 0 | 0 | 0 | 1 | 1 | 0 | 4 | 18 |
9 | *Agabus japonicus continentalis Guéorguiev, 1970 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 3 |
10 | *Agabus kaszabi Guéorguiev, 1972 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 1 | 1 |
11 | Agabus lapponicus Thomson, 1867 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 6 |
12 | *Agabus moestus Curtis, 1835 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 1 |
13 | *Agabus thomsoni Sahlberg, 1871 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 2 |
14 | Agabus clavicornis Sharp, 1882 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 2 | 9 |
15 | Agabus lineatus Gebler, 1848 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 1 |
16 | Agabus pallens Poppius, 1905 | 1 | 0 | 1 | 0 | 0 | 0 | 1 | 1 | 0 | 0 | 4 | 17 |
17 | Agabus adpressus Aubé, 1837 | 1 | 1 | 1 | 1 | 0 | 1 | 0 | 1 | 1 | 0 | 7 | 58 |
18 | *Agabus biguttulus Thomson, 1867 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 1 | 1 |
19 | *Agabus blatta Jakovlev, 1897 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 2 | 2 |
20 | Agabus dichrous Sharp, 1878 | 1 | 1 | 1 | 1 | 0 | 0 | 0 | 1 | 0 | 0 | 5 | 40 |
21 | Agabus kholini Nilsson, 1994 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 1 | 1 |
22 | Agabus laferi Nilsson, 1994 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 2 |
23 | Agabus nebulosus Forster, 1771 | 0 | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 2 | 2 |
24 | *Agabus svenhedini Falkenström, 1932 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 2 | 2 |
25 | Agabus udege Nilsson, 1994 | 0 | 0 | 0 | 1 | 0 | 1 | 0 | 1 | 0 | 0 | 3 | 4 |
26 | Ilybius angustior Gyllenhal, 1808 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 2 | 12 |
27 | Ilybius balkei Fery&Nilsson, 1993 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 1 |
28 | Ilybius chishimanus Kôno, 1944 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 9 |
29 | Ilybius cinctus Sharp, 1878 | 0 | 1 | 0 | 0 | 1 | 1 | 1 | 0 | 0 | 0 | 4 | 8 |
30 | Ilybius erichsoni Gemminger & Harold, 1868 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 4 |
31 | Ilybius lateralis Gebler, 1832 | 1 | 0 | 0 | 0 | 0 | 0 | 1 | 1 | 1 | 0 | 4 | 8 |
32 | Ilybius obtusus Sharp, 1882 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 1 | 1 | 0 | 3 | 10 |
33 | Ilybius opacus Aubé, 1837 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 2 |
34 | Ilybius poppiusi Zaitzev, 1907 | 1 | 1 | 0 | 0 | 1 | 0 | 1 | 1 | 0 | 0 | 5 | 26 |
35 | Ilybius subaeneus Erichson, 1837 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 2 | 5 |
36 | Platambus maculatus Linnaeus, 1753 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 1 |
Colymbetinae | |||||||||||||
37 | Colymbetes dahuricus Aubé, 1837 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 1 | 0 | 0 | 2 | 8 |
38 | *Colymbetes dolabratus Paykull, 1798 | 1 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 2 | 4 |
39 | Colymbetes pseudostriatus * Nilsson, 2002 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 1 |
40 | Rhantus frontalis Marsham, 1802 | 1 | 1 | 1 | 1 | 1 | 0 | 1 | 1 | 0 | 0 | 7 | 16 |
41 | Rhantus notaticollis Aubé, 1837 | 1 | 0 | 1 | 0 | 0 | 1 | 1 | 1 | 1 | 0 | 6 | 62 |
42 | *Rhantus rufus Zimmermann, 1922 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 1 | 1 |
43 | Rhantus vermiculatus Moschulsky, 1860 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 2 | 0 | 2 | 12 |
Dytiscinae | |||||||||||||
44 | Acilius canaliculatus Nicolai, 1822 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 1 | 0 | 2 | 2 |
45 | *Acilius sulcatus Linnaeus, 1758 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 1 |
46 | Graphoderus austriacus Sturm, 1834 | 1 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 1 | 0 | 4 | 4 |
47 | Graphoderus cinereus Linnaeus, 1758 | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 3 | 3 |
48 | Graphoderus zonatus verrucifer Sahlberg, 1824 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 1 |
49 | Graphoderus zonatus zonatus Hoppe, 1795 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 1 | 0 | 3 | 3 |
50 | *Dytiscus circumcinctus Ahrens, 1811 | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 3 | 3 |
51 | Dytiscus dauricus Gebler,1832 | 1 | 1 | 0 | 1 | 0 | 1 | 1 | 1 | 0 | 0 | 6 | 6 |
52 | Dytiscus latro Sharp, 1882 | 1 | 0 | 0 | 1 | 0 | 0 | 0 | 1 | 0 | 0 | 3 | 3 |
53 | *Hydaticus aruspex Clark, 1864 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 1 |
54 | *Hydaticus continentalis Balfour-Browne, 1944 |
0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 1 |
Hydroporinae | |||||||||||||
55 | *Bidessus nasutus Sharp, 1887 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 2 | 8 |
56 | *Bidessus unistriatus Goeze, 1777 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 1 |
57 | Hydroglyphus geminus Fabricius, 1792 | 0 | 1 | 1 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 3 | 16 |
58 | *Hydroglyphus hamulatus Gyllenhal, 1813 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 1 |
59 | Hydroglyphus licenti Feng, 1936 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 1 |
60 | *Boreonectes aff. emmerichi Falkenström, 1936 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 3 |
61 | Nebrioporus airumlus Kolenati, 1845 | 1 | 1 | 1 | 1 | 0 | 0 | 0 | 1 | 1 | 0 | 6 | 32 |
62 | *Nebrioporus assimilis Paykull, 1798 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 1 |
63 | Nebrioporus depressus Fabricius, 1775 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 3 | 4 |
64 | *Nebrioporus hostilis Sharp, 1884 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 2 | 3 |
65 | Oreodytes mongolicus Brinck, 1943 | 1 | 1 | 1 | 1 | 0 | 1 | 0 | 1 | 1 | 1 | 7 | 40 |
66 | Oreodytes septentrionalis Gyllenhal in C.R. Sahlberg, 1826 | 1 | 0 | 1 | 1 | 0 | 1 | 0 | 1 | 0 | 1 | 5 | 27 |
67 | Oreodytes shorti Shaverdo & Fery, 2006 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 2 | 4 |
68 | Nectoporus sanmarkii Sahlberg, 1826 | 1 | 1 | 1 | 1 | 0 | 0 | 0 | 1 | 0 | 0 | 5 | 45 |
69 | Zaitzevhydrus formaster Zaitzev, 1908 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 1 | 0 | 3 | 20 |
70 | Hydroporus acutangulus complex Thomson, 1856 | 1 | 0 | 1 | 0 | 0 | 1 | 1 | 1 | 1 | 0 | 6 | 36 |
71 | Hydroporus angusi Nilsson, 1990 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 1 |
72 | *Hydroporus crinitisternus Shaverdo & | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 1 |
73 | Hydroporus elongatulus Sturm, 1835 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 2 |
74 | Hydroporus fuscipennis Schaum & Kiesenwetter, 1867 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 3 |
75 | Hydroporus geniculatus Thomson, 1856 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 1 | 0 | 3 | 9 |
76 | Hydroporus kabakovi Fery & Petrov, 2006 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 2 |
77 | Hydroporus morio Aubé, 1838 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 5 |
78 | Hydroporus nigellus Mannerheim, 1853 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 3 |
79 | Hydroporus notabilis LeConte, 1850 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 6 |
80 | Hydroporus palustris Linnaeus, 1760 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 1 |
81 | Hydroporus sibiricus Sahlberg, 1880 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 1 | 0 | 0 | 2 | 5 |
82 | Hydroporus submuticus Thomson, 1874 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 9 |
83 | Hydroporus uenoi Nakane, 1963 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 13 |
84 | Hygrotus caspius Wehncke, 1875 | 0 | 0 | 0 | 0 | 1 | 1 | 0 | 1 | 0 | 0 | 3 | 10 |
85 | Hygrotus confluens Fabricius, 1787 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 1 |
86 | Hygrotus enneagrammus Ahrens, 1833 | 1 | 1 | 1 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 4 | 9 |
87 | Hygrotus flaviventris Motschulsky, 1860 | 1 | 1 | 1 | 1 | 1 | 1 | 0 | 1 | 0 | 1 | 7 | 24 |
88 | Hygrotus nigrolineatus Steven in Schönherr, 1808 | 1 | 1 | 0 | 0 | 1 | 0 | 0 | 1 | 0 | 0 | 4 | 6 |
89 | Hygrotus pectoralis Motschulsky, 1860 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 3 | 3 |
90 | Hygrotus inaequalis Fabricius, 1777 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 1 | 1 | 0 | 3 | 8 |
91 | Hygrotus quinquelineatus Zetterstedt, 1828 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 1 | 0 | 3 | 17 |
92 | Hygrotus chinensis Sharp, 1882 | 0 | 1 | 0 | 0 | 0 | 1 | 0 | 1 | 0 | 0 | 3 | 4 |
93 | Hygrotus impressopunctatus Schaller, 1783 | 1 | 1 | 1 | 1 | 0 | 1 | 0 | 1 | 1 | 0 | 7 | 72 |
94 | Hygrotus marklini Gyllenhal, 1813 | 1 | 1 | 0 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 9 | 53 |
95 | Hygrotus parallellogrammus Ahrens, 1812 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 1 | 0 | 0 | 2 | 3 |
96 | Hygrotus unguicularis Crotch, 1874 | 1 | 1 | 1 | 0 | 0 | 1 | 0 | 1 | 0 | 0 | 5 | 31 |
97 | Hygrotus urgensis Jakovlev, 1899 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 1 |
98 | *Laccornis oblongus Stephens, 1835 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 1 |
Laccophilinae | |||||||||||||
99 | Laccophilus biguttatus Kirby, 1837 | 1 | 1 | 0 | 1 | 0 | 1 | 1 | 1 | 1 | 0 | 7 | 44 |
Overall species composition | 45 | 26 | 23 | 21 | 9 | 17 | 16 | 79 | 21 | 5 |
Species list of Dytiscid (Coleoptera, Dytiscidae) of Mongolia
Data type: Table data