Patterns and Processes of Speciation in Ancient Lakes Developments in Hydrobiology 205 Series editor K. Martens Patterns and Processes of Speciation in Ancient Lakes Proceedings of the Fourth Symposium on Speciation in Ancient Lakes, Berlin, Germany, September 4–8, 2006 Editors Thomas Wilke1, Risto Va¨ino¨la¨2 & Frank Riedel3 1JustusLiebigUniversityGiessen,DepartmentofAnimalEcologyandSystematics,Heinrich-Buff-Ring26-32(IFZ),D-35392 Giessen,Germany 2FinnishMuseumofNaturalHistory,POBox26,FI-00014UniversityofHelsinki,Finland 3FreeUniversityofBerlin,DepartmentofEarthSciences,Malteserstr.74-100,HouseD,D-12249Berlin,Germany Reprinted from Hydrobiologia, Volume 615 (2008) 123 LibraryofCongressCataloging-in-PublicationData AC.I.P.CataloguerecordforthisbookisavailablefromtheLibraryofCongress. ISBN:978-1-4020-9581-8 PublishedbySpringer, P.O.Box17,3300AADordrecht,TheNetherlands CitethispublicationasHydrobiologiavol.615(2008). Coverillustration:EasternshoreofLakeOhridnearVeliDab–oneofthelake’smajorhotspotsof endemicbiodiversity.Photograph:ThomasWilke,2004. Printedonacid-freepaper AllRightsreserved (cid:2)2008SpringerScience+BusinessMediaB.V. Nopartofthismaterialprotectedbythiscopyrightnoticemaybereproducedorutilizedinanyform orbyanymeans,electronicormechanical,includingphotocopying,recordingorbyanyinformation storageandretrievalsystem,withoutwrittenpermissionfromthecopyrightowner. TABLE OF CONTENTS Preface T. Wilke, R. Va¨ino¨la¨, F. Riedel 1–3 EAST AFRICAN GREAT LAKES The Lake Tanganyika cichlid species assemblage: recent advances in molecular phylogenetics S. Koblmu¨ller, K.M. Sefc, C. Sturmbauer 5–20 Variance in reproductive success and the opportunity for selection in a serially monogamous species: simulations of the mating system of Tropheus (Teleostei: Cichlidae) K.M. Sefc 21–35 Assortative mating preferences between colour morphs of the endemic Lake Tanganyika cichlid genus Tropheus B. Egger, B. Obermu¨ller, E. Eigner, C. Sturmbauer, K.M. Sefc 37–48 VariationofterritorysizeanddefensebehaviorinbreedingpairsoftheendemicLake Tanganyika cichlid fish Variabilichromis moorii C. Sturmbauer, C. Hahn, S. Koblmu¨ller, L. Postl, D. Sinyinza, K.M. Sefc 49–56 Abundance,distribution,andterritoryareasofrock-dwellingLakeTanganyikacichlid fish species C. Sturmbauer, C. Fuchs, G. Harb, E. Damm, N. Duftner, M. Maderbacher, M. Koch, S. Koblmu¨ller 57–68 Subtlepopulationstructureandmale-biaseddispersalintwoCopadichromisspecies (Teleostei, Cichlidae) from Lake Malawi, East Africa D. Anseeuw, G.E. Maes, P. Busselen, D. Knapen, J. Snoeks, E. Verheyen 69–79 Ecological correlates of species differences in the Lake Tanganyika crab radiation S.A.E. Marijnissen, E. Michel, M. Kamermans, K. Olaya-Bosch, M. Kars, D.F.R. Cleary, E.E. van Loon, P.G. Rachello Dolmen, S.B.J. Menken 81–94 The Great Lakes in East Africa: biological conservation considerations for species flocks C. Sturmbauer 95–101 BALKAN LAKES OHRID AND PRESPA Ancient Lake Ohrid: biodiversity and evolution C. Albrecht, T. Wilke 103–140 Theneglectedsideofspeciationinancientlakes:phylogeographyofaninconspicuous mollusctaxoninlakesOhridandPrespa R. Schultheiß, C. Albrecht, U. Bo¨ßneck, T. Wilke 141–156 vi Concurrent evolution of ancient sister lakes and sister species: the freshwater gastropod genus Radix in lakes Ohrid and Prespa C. Albrecht, C. Wolff, P. Glo¨er, T. Wilke 157–167 Testing two contrasting evolutionary patterns in ancient lakes: species flock versus species scatter in valvatid gastropods of Lake Ohrid A.-K. Hauswald, C. Albrecht, T. Wilke 169–179 SULAWESI LAKE SYSTEMS The species flocks of lacustrine gastropods: Tylomelania on Sulawesi as models in speciation and adaptive radiation M. Glaubrecht, T. von Rintelen 181–199 Gene flow at the margin of Lake Matano’s adaptive sailfin silverside radiation: Telmatherinidae of River Petea in Sulawesi J. Schwarzer, F. Herder, B. Misof, R.K. Hadiaty, U.K. Schliewen 201–213 Processes regulating the community composition and relative abundance of taxa in the diatom communities of the Malili Lakes, Sulawesi Island, Indonesia A.J. Bramburger, P.B. Hamilton, P.E. Hehanussa, G.D. Haffner 215–224 The plankton community of Lake Matano: factors regulating plankton composition and relative abundance in an ancient, tropical lake of Indonesia E. Sabo, D. Roy, P.B. Hamilton, P.E. Hehanussa, R. McNeely, G.D. Haffner 225–235 Hydrobiologia(2008)615:1–3 DOI10.1007/s10750-008-9559-x SPECIATION IN ANCIENT LAKES Preface Thomas Wilke Æ Risto Va¨ino¨la¨ Æ Frank Riedel (cid:2)SpringerScience+BusinessMediaB.V.2008 (cid:3) Ancientlakes,thatis,extantlakesthattypicallyhave lakes. Thomson Reuter’s Web of Science , for continuouslyexistedsincebeforethePleistocene,for example, lists under this topic, only one cited paper more than two million years—such as lakes Baikal, in 1996, but 190 for the year 2007. Tanganyika, Malawi, Biwa, and Ohrid, have long This volume represents the outcome of a fourth beenrecognizedascentresofbiodiversity.Moreover, meeting in a series of conferences on speciation in during the past years, speciation in ancient lakes has ancient lakes (SIAL). The series was started by a emerged as an important and fruitful topic in studies meeting that took place in 1993 in Mont-Rigi, ofevolutionarybiology.Thecurrentunderstandingof Belgium, and was brought together to review the a number of key evolutionary concepts, such as state of art in the field for the first time since the sympatric speciation, sexual selection, adaptive radi- classical seminal review of Brooks (1950). The ation,hybridization,andpunctuatedequilibrium,isin proceedings of that meeting were published as a partbasedoninsightsfromspeciationstudiesinthese specialvolumeofArchivfu¨rHydrobiologie(Martens lakes, and this is evident in the increasing number of et al.,1994);apartfromsummarizingboththeoretical citationsofarticlespertainingtospeciationinancient concepts of evolution in ancient lakes and practical activitiesinthesesystems,italsoinitiatedawealthof renewed interest in ancient lake issues. The second SIAL meeting entitled ‘‘International Guesteditors:T.Wilke,R.Va¨ino¨la¨ &F.Riedel Conference on Ancient Lakes: their Biological and PatternsandProcessesofSpeciationinAncientLakes: ProceedingsoftheFourthSymposiumonSpeciationin Cultural Diversities’’ was held in 1997 near Lake AncientLakes,Berlin,Germany,September4–8,2006 Biwa in Japan. It focused on the cultural diversity of ancient lakes (Kawanabe et al., 1999) as well as on T.Wilke(&) theirbiodiversity,ecology,andevolution(Rossiter& DepartmentofAnimalEcologyandSystematics, JustusLiebigUniversityGiessen,Heinrich-Buff-Ring Kawanabe, 2000). 26-32(IFZ),35392Giessen,Germany The third conference, SIAL-3, took place in e-mail:[email protected] Irkutsk. Naturally, the focus of that conference was the oldest and arguably most famous of the ancient R.Va¨ino¨la¨ FinnishMuseumofNaturalHistory,Universityof lakes—Lake Baikal. The conference proceedings Helsinki,POBox17,00014Helsinki,Finland were published in two special issues (Ivanov et al., 2003; Coulter et al., 2006), adding to the reputation F.Riedel of Baikal as one of the best-studied lakes in the DepartmentofEarthSciences,FreeUniversityofBerlin, Malteserstr.74-100,HouseD,12249Berlin,Germany world. 123 2 Hydrobiologia(2008)615:1–3 Finally, the latest meeting SIAL-4 was held on strategies of cichlid fish species in Lake Tanganyika September4–8,2006,attheFreieUniversita¨tBerlin, (Sefc, Egger et al. and Sturmbauer et al.) and Lake Germany, organized by Frank Riedel (Germany) Malawi (Anseeuw et al.). An endemic radiation of togetherwith George Coulter (New Zealand),Matth- freshwatercrabsinLakeTanganyikaisthesubjectof ias Glaubrecht (Germany), Doug Haffner (Canada), the study of Marijnissen et al., who discuss data on Hiroya Kawanabe (Japan), Koen Martens (Belgium), interspecific spatial and ecological segregation in the Oleg Timoshkin (Russia), Risto Va¨ino¨la¨ (Finland), framework of adaptive radiation theory. Finally, and Thomas Wilke (Germany); 67 scientists from 16 Sturmbauerreviewsanddiscussesconservationstrat- different countries contributed to the scientific pro- egies for aquatic biota and their biological impact on gram. While the abstracts of SIAL-4 were published species flocks in the East African Great Lakes. in a special issue of Berliner Pala¨obiologische Abhandlungen (Kossler et al., 2006), selected full papers are subject of the present special issue The Balkan lakes Ohrid and Prespa ‘‘Patterns and processes of Speciation in Ancient Lakes.’’ The contributions in this issue deal with OneofthemostoutstandingEuropeanancientlakesis threeancientlakessystems.Onesystemisthefamous theoligotrophicandkarsticLakeOhridsituatedinthe East African Great Lakes, which have been studied central Balkans. Together with its sister lake, Lake extensively over the past decades, resulting in Prespa, it represents one of the most significant significantnewfindingsaboutevolutionaryprocesses hotspots of endemic biodiversity in the world. in general. On the other hand, we include a set of AlthoughtherecognitionofLakeOhridasapeculiar papersontwootherprominentlakesystemsthathave lakewithanoutstandingbiodiversitybeganinthelate so far received much less attention, the European 19thcentury,andbasictaxonomicalandevolutionary ancient sister lakes Ohrid and Prespa, which have studieswerecarriedoutpriortoandafterWorldWar long been recognized as hotspots of biodiversity but II,thepoliticalinstabilitiesintheBalkansinthe1980s were largely neglected in the international scientific and1990sledtoasharpdeclineofprojectscarriedout literature until recently, and the two central lake attheselakes.Onlythelastdecadehasseenarevival systemsontheIndonesianislandofSulawesi,therich ofscientific activity inlakesOhrid andPrespa. biodiversity and interesting evolutionary patterns of Here, Albrecht and Wilke summarize the current which only very recently have started to draw the knowledge of the limnological and faunal history of attention of evolutionary biologists. Lake Ohrid, update the existing data on its biodiver- sityandendemism,andreviewpatternsandprocesses ofspeciationinthelake.Inprimary research studies, East African Great Lakes several groups of endemic molluscs are investigated with regard to the sister-lake relationship of lakes TheAfricanGreatLakes—aseriesoflakesinornear Ohrid and Prespa (Albrecht et al.), patterns of theGreatRiftValley,forexampleLakeVictoria,Lake endemic biodiversity (Hauswald et al.), and evolu- Tanganyika,andLakeMalawi—aremajorhotspotsof tionary patterns in inconspicuous (cryptic) taxa freshwater biodiversity, with splendid examples of (Schultheiß et al.). large radiations of cichlid fishes. While notall of the African Great Lakes qualify as ancient (e.g., the relativelyyoungLakeVictoria),theselakeshavelong Sulawesi lake systems beena‘‘playground’’forevolutionarybiologists. In the current issue, Koblmu¨ller et al. give an While evolutionary studies in ancient lakes such as accountofthehighertaxonomyandphylogenyofthe LakeBaikalandLakeTanganyikahavebeenongoing Lake Tanganyika cichlid species assemblage, its for many decades, patterns of endemic diversity in relationship to the African cichlid fauna, and key thetwocentrallakesystemsontheIndonesianisland factorsleadingtotheastonishingdiversityofcichlids of Sulawesi, that is, Lake Poso and the lakes of the in the lake. Several other workers report on the Malili lakesystem,have been largelyneglected fora emerging field of sexual selection and reproductive century.Recently,however,therehasbeenarenewed 123 Hydrobiologia(2008)615:1–3 3 interest in these lakes and in their endemic species topics will certainly continue to be focal points flocks. during the fifth conference, SIAL-5, which is sched- In the present issue, Glaubrecht and Rintelen uled to take place at Lake Ohrid in September 2009. review and discuss the evolutionary and taxonomic We would like to thank the other members of the implications of the Tylomelania gastropod species organizing committee for their hard work. We also flocks and demonstrate that they provide instructive would like to thank Koen Martens, who kindly gave model cases for the study of speciation mechanisms, us the opportunity to publish this issue and who adaptiveradiation,andnicheexploitation.Schwarzer assisted us during preparation. Eugene Coan is et al. utilized the adaptive radiation of sailfin silver- gratefully acknowledged for improving a number of sides in Lake Matano (Malili lake system) to study manuscriptsasanativeEnglishspeakerandbiologist, the effect of gene flow on speciation processes. The and Claudia Wilke helped with proofreading. datapresentedsuggestthatgeneticexchangebetween The Guest Editors lacustrine and riverine taxa takes place, potentially allowing introgression into Lake Matano from lower lakes and streams. The rich endemic diatom flora of the Malili lake system is the subject of the study of References Bramburger et al. The workers reviewed the mech- anisms contributing to the development and Brooks, J. L., 1950. Speciation in ancient lakes. Quarterly ReviewofBiology25(30–60):131–176. maintenance of community composition and relative Coulter,G.,O.Timoshkin,L.Timoshkina&K.Martens(eds), abundance of taxa. Finally, Sabo et al. studied the 2006.Speciesandspeciation inancientlakes.Hydrobio- phytoplanktonandzooplanktoncommunitiesofLake logia568(S):287pp. Matano and substantiated the assumption that Ivanov,A.,G.Coulter,O.Timoshkin&F.Riedel(eds),2003. Proceedings of the international symposium speciation resourcelimitationplaysanimportantroleinshaping in ancient lakes, SIAL III. Berliner Pala¨obiologische the unique endemic assemblages currently observed Abhandlungen4:200pp. in the food web of the lake. Kawanabe,H.,G.W.Coulter&A.C.Roosevelt(eds),1999. We hope that the present special issue can deepen Ancient lakes: their cultural and biological diversity. KenobiProductions,Ghent,Belgium. the awareness of ancient lakes in the scientific Kossler,A.,R.Kohring&F.Riedel(eds),2006.Abstractsof community in general, and of their role as natural the international symposium speciation in ancient lakes, laboratories and prime hotspots of biodiversity in SIAL IV. Berliner Pala¨obiologische Abhandlungen 9: particular. It is also hoped that this issue stimulates 70pp. MartensK.,B.Goddeeris&G.Coulter(eds),1994.Speciation furtherresearchthatmayleadtoabetterunderstand- inancientlakes.AdvancesinLimnology44:508pp. ing of key evolutionary processes in these lakes and Rossiter, A. & H. Kawanabe (eds), 2000. Ancient lakes: bio- that ultimately may help in mitigating the decline of diversity,ecologyandevolution.AdvancesinEcological biodiversity seen today in many ancient lakes. These Research31:624pp. 123
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