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Diversity and Conservation of the Northern Far East Asian Freshwater Fishes, with a Case Study on Freshwater Sculpins

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Diversity and Conservation of the Northern Far East Asian Freshwater Fishes, with a Case Study on Freshwater Sculpins 57 Diversity and conservation of the northern Far East Asian freshwater fishes, with a case study on freshwater sculpins AKIRA GOTO GraduateSchool of Fisheries Sciences,Hokkaido University,Hakodate 041-8611,Japan ([email protected]) SUMMARY: The freshwater fishes in the northern Far East were investigated to elucidate the species richness, ecological complexity and genetic diversity. The fishes in Hokkaido Island, Sakhalin Island, Primor'e and the Amur River basin were composed of 27 families, 83 genera and 130 species which possessed high ecological and genetic diversity. Concerning the freshwater sculpins, for example, one Trachidermus, one Mesocottus and 11 Coitus species are distributed in the regions. The mitochondrial DNA analyses made clear that the amphidromous, lacusbine and fluvial species have originated from the ancestral stocks of catadromous species such as T fasciatus and Coitus kazika, showing the creation of species diversity. Mainly because of rapid changes of freshwater environments, however, nearly 10-40 % fish species to the total number of species distributed in the regions were found to be endangered, indicating the necessity of effective conservation actions. KEY WORDS: species diversity, life-history diversity, genetic variability, phylogenetic relationships, speciation, endangered fish INTRODUCTION Therefore, it is reasonable to state that we should face The freshwatersare composedof rivers, streams, lakes, the research subjects of freshwater fish diversity and their swamps,waterways in rice fieldand so on, in which fish conservation in both global and local scales. The main specieshaving various modes of life-historieslive: those are subjects for understandingthe diversity of freshwater fishes fluvial, anadromous, catadromous, amphidromous, in the northern Far East are as follows: the first is to estuarine,lacustrine and so forth. The diversityof these elucidate the species richness which includes abundance of freshwaterfishes are characterizedby hierarchicalstructures species numbers and population sizes in each species.; the from genes to ecosystems through populations and second is to make clear the ecological diversity in which communities(Fig. 1). 1) complexity of life-history patterns and reproductive styles, In the last decade, however, the environmentsof population dynamics and interspecificrelationships in fishes feshwatershave been rapidlychanged mainly by great are included; and the third is to elucidatethe variability of increaseof human activitieswhich had resultedin global genetic population structure and phylogeography in each warming,water pollution, reclamation of streamsand lakes, fish species, and phylogenetic relationships among the artificialreservoirs and so on.23)As a result, freshwater species. fishes had heavy damages on number of species and populationsizes. DIVERSITY AND CONSERVATION OF THE For example,an estimationof habitat losses in cold NORTHERN FAR EAST ASIAN FRESHWATER - water adapted fish by globally climaticwarming was FISHES carriedout using two chars, Salvelinusmalma and S. leucomaenis,and demonstratedthat the habitat losses of In order to estimate the diversity of freshwater fishes in chars will advanceaccording to the increaseof average northern Far East, our research group have had several annualtemperature.4) expeditionsin the regions for recent 10 years. In this paper, I describebriefly about some findings of species, ecological 58 Fig. 1. Habitat diversity and hierarchical structure of biodiversity in the freshwater fishes. S: spawning, G: growth. and genetic diversitiesof fishes. amphidromous fish species in Hokkaido Island which are The expeditions were carried out mainly in Hokkaido composed of Plecoglossus altivelis,3 Cottus species and 12 Island, Japan and Sakhalin Island, Primor'e and the Amur gobiid species.) This may be due to the past northward River basin in Russia Concerning the species richness, the dispersalof these fishesfrom Honshu Island. native freshwater fishes are composed of 27 families, 83 About the diversity of reproductive styles, only the genera and about 130 species in all the four regions. In each feature in Hokkaido Island is presented because of almost region, Hokkaido is inhabited by 36 genera 59 species, no data in Russia Hokkaido's freshwater fishes are Sakhalin by 39 genera 39 genera 57 species, Primor'e by 33 limited to have 7 reproductivestyles. As nonguarders,there genera 45 species, and Amur River basin by 70 genera 90 are 4 styles, and the other 3 styles are found in guarders;For species, demonstratingthe most species richness in Amur example, osmerid species are nonguarders and have the River basin (Table 1).5) style as rock and gravel spawners with pelagic larvae. On From the view point of life-historytypes, these fishes are the other hand, gobiid species are guarders and have the classified into pure fieshwater fish, secondary freshwater styles as hole nesters.7) fish, diadromous fish and peripheral fish. Furthermore, Concerning the genetic diversity, we have obtained diadromous fishes are subdivided into anadromous, some new findings of peculiar genetic structure and genetic catadromous and amphidromous fish 6,7)Comparing to the divergence in several taxa of the norther Far Eastern poorness of pure freshwater fish species in Hokkaido, freshwater fishes. Of the findings, following two examples Sakhalin and Primor'e, remarkably abundant species are are shown: One is for threespine sticklebacks and the other found in Amur River basin. The species richness of these is for Japanese fluvialsculpins. fishes in Amur River basin is supposed to have occurred In threespine sticklebacks,it has been believed that they mainly due to the connection of this river with a branch of comprise only one species, Gasterosteus aculeatus around Yellow River basin about 100 thousands years ago.8,99In Japan. Our recent studies on the fishes, however, addition, one more peculiarityis found in the abundance of demonstratedclearly that two high-geneticallydivergent 59 Table 1. Abundance of fish species distributed in freshwaters of abundance of extinct and endangeredspecies among several the northern Far East regions or countries shows that the percentage to the total number of species is considerably high in whole Japan, whereas that in Hokkaido Island is not so high (Table 2). The official conservationactions should be performed for the endangeredfreshwater species in Japan. Table 2. Comparison of abundance of endangered freshwater fish species among several regions A CASE STUDY ON THE ORIGIN OF SPECIES DIVERSITY IN FRESHWATER SCULPINS In the Far Eastern Asia, the freshwater sculpins are Figures in parentheses indicate the number of genera. composed of one Trachidermus, one Mesocottus and 11 forms(Nei's D= 0.43) exist around Japan.10)One form was Cottus species, which have various types of life-histories convenientlynamed as Pacific Ocean foam,and the other as such as ratadromous, amphidromous, lacustrine and fluvial Japan Sea form. The two forms were found to be types. reproductivelyisolated from each other in several sympatric In Japan, for example, the freshwater sculpins are populations.Therefore, the two forms should be classified thought to comprise one Trachidermus and 7 Cottus as differentspecies. The other example is of a cold-water species.14)Of the species, Trachidermus fasciatus and adaptedsculpin, Coitus nozawae. This species is endemic to Coitus kazika are catadromous, C. amblystomopsis, C Japan,distributed in Hokkaido Island and Tohoku District, pollwr SE, C pollux ME and C hangiongensts arty the northernmostpart of Honshu Island. The mitochondrial amphidromous,C. reinii, which are endemic to Lake Biwa, DNA analysisshowed a great genetic divergence between is lacustrine,and C nozawae and C. pollux LE are fluvial. two regional populations from Hokkaido and Tohoku Recent allozyme and mtDNA studies showed that Japanese District.11) In the southern margin of its range, moreover, freshwater sculpin species comprise two major clades and haplotype displacementswere observed between tributary the lacustine C. reinii is a genetic variant within the populationswithin a single river-system,probably because amphidromous C. pollux SE, the latter suggesting that they of their habitatfiagmentalion in this region and their strong should be a single species (Fig. 2). Therefore, the valid sedentarrity.12) Coitus species in Japan is thought to be 7 species, and In the last two decades, however, the species diversity among them 5 species (kazika,pollux ME, pollux LE, reinii has graduallydecreased mainly because of human induced and nozawae) are endemicto Japan. environmental changes. Of about 250 freshwater and In order to elucidatethe origin, dispersal and speciation blackishwater fish species distributedin Japan, 3 species or events of these sculpins, the phylogenetic relationships subspecies were already extinct, and 88 species or among the freshwatersculpins in the Far Eastern Asia were subspeciesare now endangered(the EnvironmentalAgency, inferred from nucleotide sequences of mitochondrial 12S Japan Government).13) A comparison of the relative rRNA gene and controlregion (1,895 base-pairs). 60 are not monophyleticbut paraphyletic,because C kazikais more closelyrelated to T fasciatus than the other Coitus speciesexamined. In Group 11,several monophyletic species pairs were found Interestingly,such sister relationships supported by high bootstrapprobabilities (above 90 %) are seen between amphidromousC. amblystomopsisand fluvialC.
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