on the brink - investigating biodiversity in endangered crater lakes of the amber mountains national...

On the brink ndash investigating biodiversity in endangered crater lakesof the Amber Mountains National Park (Madagascar)

R SCHABETSBERGERa RKAISERa EROTTb R LENZENWEGERcWTRAUNSPURGERd AAKOTOVe F FIERSfS GAVIRIAg C MEISCHh S MAHUNKAi R D CAMPBELLj E RAZAFINDRANAIVOk and M SZTATECSNYl

aDepartment of Organismic Biology University of Salzburg Salzburg AustriabInstitute of Botany University of Innsbruck Innsbruck Austria

cSchlossberg 16 4910Ried im Innkreis AustriadAnimal Ecology University of Bielefeld Bielefeld Germany

eAN Severtsov Institute of Ecology and Evolution Moscow RussiafRoyal Belgian Institute of Natural Sciences Brussels Belgium

gDepartment of Limnology University of Vienna Vienna AustriahNational Museum of Natural History Luxembourg Luxembourg

iHungarian Natural History Museum Budapest HungaryjDepartment of Developmental and Cell Biology University of California Irvine USA

kDepartment of Animal Biology University of Antananarivo Antananarivo MadagascarlDepartment of Evolutionary Biology University of Vienna Vienna Austria

ABSTRACT

1 Madagascarrsquos biodiversity is vanishing at an alarming pace The documentation of this loss has so far focusedon terrestrial habitats and organisms Eight volcanic crater lakes in the AmberMountains National Park (NorthernMadagascar) and surroundings were investigated for the first time to describe limnological conditions and aquaticbiodiversity Seven of the lakes were affected by deforestationlogging and fish introduction and only one lake wasassumed to have remained in pristine condition In the deeper lakes (gt 5m) steep physico-chemical gradients andanoxic hypolimnia were observed

2 Algae hydrozoans nematodes rotifers annelids copepods cladocerans ostracods and mites were identifiedto genus or species level The majority were taxa with a cosmopolitan or tropicopolitan distribution The highestnumber of afrotropical and endemic species were recorded within the crustaceans

3 Multivariate analysis of species communities revealed significant differences between lakes in deforested andforested catchments Introduced alien fish had no detectable effect on species assemblages

4 Illegal harvest of timber was observed within the National Park and drug plantations are less than 1 km awayfrom the last pristine crater lake If deforestation continues at the current rate which is likely under the prevailingpolitical situation the last undisturbed lake communities may be altered in the near future

5 There is an urgent need for taxonomic research to assess the biodiversity of algae and micrometazoa Highestpriority should be given to pristine freshwater ecosystems within protected areasCopyright 2012 John Wiley amp Sons Ltd

Received 6 April 2012 Revised 5 July 2012 Accepted 4 August 2012

KEY WORDS algae alien species biogeography deforestation invertebrates island lake taxonomy

Correspondence to Robert Schabetsberger Department of Organismic Biology University of Salzburg Hellbrunnerstrasse 34 A-5020 SalzburgAustria Email RobertSchabetsbergersbgacat

Copyright 2012 John Wiley amp Sons Ltd

AQUATIC CONSERVATION MARINE AND FRESHWATER ECOSYSTEMS

Aquatic Conserv Mar Freshw Ecosyst 23 316ndash331 (2013)

Published online 13 November 2012 in Wiley Online Library(wileyonlinelibrarycom) DOI 101002aqc2286

INTRODUCTION

The island of Madagascar is among the tophotspots for biodiversity conservation worldwide(Myers et al 2000 Ganzhorn et al 2008 Vieiteset al 2009) More than 90 of all plants 100 ofnative amphibian and mammal species 92 ofreptiles and 44 of birds occur nowhere else Thishigh level of endemism predominantly results fromradiation of African founder individuals arrivingduring the Cenozoic (655Ma to present) and fromrelict species radiating after the island separatedfrom the African mainland (183ndash158Ma)Antarctica (130Ma) and India (96ndash65Ma Venceset al 2009)

Besides an extraordinary diversity in terrestrialorganisms Madagascar has also been recognizedas a global hotspot of freshwater biodiversity(Groombridge and Jenkins 1998) In the last twodecades of the 20th century the number of newlydescribed species of aquatic insects and fish hadincreased exponentially (Benstead et al 2003)However owing to incomplete species inventoriesand taxonomic difficulties the degree of endemismin small planktonic and benthic organisms isdifficult to assess There is evidence for endemismespecially for copepods (Dussart 1982) but alsofor some freshwater algae (eg desmids (West andWest 1895 Bourrelly and Couteacute 1991 Coesel2002) and benthic diatoms (Metzeltin and Lange ndash

Bertalot 2002))Taxonomic research on Malagasy freshwater

algae and micrometazoa is still scarce There areno systematic species inventories available andcurrent knowledge is based on random samplesmostly covering specific groups of organisms fromdifferent habitats First surveys of the Malagasyfreshwater algal flora were provided by West andWest (1895) Fritsch (1914) Manguin (1941)Bourrelly and Leboime (1946) and Bourrelly andManguin (1949) More detailed algologicalinvestigations are available for diatoms (Spauldingand Kociolek 1998a b Metzeltin and Lange ndash

Bertalot 2002) desmids (Bourrelly and Couteacute1991 Coesel 2002) and chrysophytes (Hansen1996) Taxonomic research on micrometazoa wasconducted predominantly until the 1960s(DeGuerne and Richard 1893 Brehm 19301948 Kiefer 1930) Most studies were based onsamples taken by the director of the Institut desrecherches scientifiques de Madagascar RenauldPaulian who sent the material to varioustaxonomists (Rotifera Berzins 1960 1973 1982

Harpacticoida Chappuis 1952 1954 1956Calanoida Brehm 1951 1952a b c 1953 1954Cyclopoida Lindberg 1951a b 1952 1953 Kiefer1952 1954 1955 Cladocera Brehm 1953) Brehm(1960) gave an account of freshwater Crustaceacollected during the Austrian Madagascarexpedition in 1958 After this period of intensivetaxonomic work on freshwater micrometazoa fewpublications appeared (Dussart 1982 Segers 1992Fiers 2002)

Madagascarrsquos biological richness is threatened bymassive habitat destruction Forty per cent ofrainforest cover was lost between 1950 and 2000(Harper et al 2007 Wikipedia 2012)Deforestation currently continues at a rate ofabout 05 per year (57 000 ha Mongabay 2011)Although the country possesses 46 legallyprotected areas covering almost 17 millionhectares (289 of total territory) habitatdestruction still proceeds at an alarming pace Apeak in illegal logging was observed during thepolitical turmoil in 2009 Schuurman and Lowry(2009) spoke of a lsquoMadagascar rosewoodmassacrersquo because 625 containers of hardwoodworth approximately US $ 130 million illegally leftthe country in 2009 Most of the trees were felledin protected mountain reserves in NorthernMadagascar (Marojeji Masoala and AmberMountains) During sampling in May 2009 theillegal harvest of rosewood and palisander(Dalbergia spp) was witnessed on several occasions

Deforestation not only changes terrestrialhabitats but causes increased sediment deliveryand nutrient input into rivers and lakes and hencealters physico-chemical conditions speciescomposition and trophic interactions in aquatichabitats as well (Benstead et al 2003) Loss offorest cover has already caused a shift frompersistent to intermittent flow in the streamsdraining the northern and eastern slopes of theAmber Mountains and the Tsaratanana Massif(Raxworthy et al 2008)

Following habitat destruction invasive speciesare regarded as the second leading cause of speciesextinction and endangerment worldwide (Salaet al 2000) A number of alien fish species havebeen stocked in pristine Malagasy fresh waterscausing severe changes to entire ecosystems andbeing detrimental to various groups of animals(Canonico et al 2005) For instance there is astrong correlation between the introduction ofexotic fish and the decline of native fish inMadagascar (Reinthal and Stiassny 1991 Sparks

MADAGASCAR FRESHWATER BIODIVERSITY 317

Copyright 2012 John Wiley amp Sons Ltd Aquatic Conserv Mar Freshw Ecosyst 23 316ndash331 (2013)

and Stiassny 2003) but no information is availableabout their impact on plankton and benthoscommunities in Malagasy lake ecosystems

Finally the warming trends reported forMadagascar equal or exceed global averages(Raxworthy et al 2008) A drier climate willaccelerate the destruction of montane rainforestsin the Amber Mountains National Park At itslower level the thinning of the protective canopyexposes the forest to sunlight and drying windthereby increasing the risk of wildfires

Benstead et al (2003) defined three majorobjectives to conserve the remaining Malagasyfreshwater biodiversity (1) Survey efforts have tobe directed at remote regions that have noinventories for freshwater biota (2) Systematicand ecological studies of poorly known taxonomicgroups must be undertaken (3) Top conservationpriority should be given to intact freshwaterecosystems that are situated within protected areas

Today only a few pristine volcanic crater lakeson the entire island remain surrounded by primaryrainforest (Schabetsberger et al 2009a) The twolargest clusters of crater lakes are within or nearthe borders of the volcanic massifs of the AmberMountains and the Tsaratanana (Bemanevikaarea) National Parks in Northern MadagascarThe Bemanevika lakes were found to be the lastrefuge for only a few individuals of theMadagascar pochard a duck proclaimed extinctand rediscovered in 2006 (TWSG News 2006)but the catchment already exhibits fragmentedpockets of rainforest near the lakes A singleisolated crater lake in the Makira protected areaLake Amparihibe also appeared to be pristine(Schabetsberger et al 2009a) It was reached by a1-day boat journey and a 3-day march throughrainforest Nevertheless the illegal harvest ofhardwood trees was witnessed more than half wayalong the trail These last examples of undisturbedlentic species communities which have persistedbefore and since the arrival of man onMadagascar approximately 2300 years ago(Burney et al 2004) should therefore be givenhigh priority in future conservation measures

The aim of this study was a preliminarylimnological characterization of lakes in the AmberMountains National Park and its surroundingsPreliminary species lists of planktonic and benthicorganisms are presented and the impact ofdeforestation and stocking with alien fish specieson freshwater communities is analysed Thesamples were collected during a single expedition

mounted during times of political unrest andincreasing illegal timber harvest within the AmberMountains National Park (Schuurman and Lowry2009 personal observations) Although we wereunable to collect replicate samples and measurenutrient chemistry these first baseline data areprovided to support future conservation efforts

STUDY AREA

The Amber Mountains are a large stratovulcano innorthern Madagascar (c 2500 km2) Radiometricages of lavas range from 121 02Ma (40Ar-39Ar)in the north west to 083 002Ma (40K40Ar) inthe central part suggesting that the volcanism wasactive from at least the middle Miocene Themassif is formed by lsquohundreds of lava flows plugsspatter cones tuff rings pyroclastic flows andpyroclastic fall depositsrsquo (Cucciniello et al 2011and references therein) Several lakes fill tuff ringsthat represent the youngest volcanic activity of thecomplex

In 1958 the FrenchColonial Forces established thefirst Malagasy National Park in the AmberMountains massif Today the Park protects18 200 ha of mid-altitude montane rainforest lyingat altitudes ranging from 850ndash1475m Five volcaniccrater lakes are situated within (here numbered(1ndash3) (7)) or at the edge of the forest (8) whilethree lakes at lower altitude (4ndash6) are outside thepark and are surrounded by savannah (Figures 1 2Table 1) Lake Manonja (7) remains the onlyunstocked lake surrounded by primary rainforestwhereas the savannah has already reached thesouthern shore of Lake Fantany (8) Also lsquokhatrsquo(Catha edulis ndash a drug common in East Africa)plantations are closing in on Lake Manonja and arenow less than 1 km away

During French occupation a forest stationoperated at the park entrance near Joffreville(A Randimbison personal communication) Fromthere black bass (Micropterus salmoides) and carp(Cyprinus carpio) were introduced into three out offive lakes in the centre of the park (Mahasarika(1) Malio (2) Taranta (3)) In addition tilapia(Tilapia rendalli) and mosquitofish (Gambusiaaffinis) are found today in the lake closest to thepark entrance (Mahasarika (1)) The lakessurrounded by savannah (Mahery (4) Antagnavo(6) and Antagnavo ely (5)) contain tilapiastogether with endemic cichlids and are inhabitedby crocodiles (Crocodylus niloticus)

R SCHABETSBERGER ET AL318


METHODS

Field work lasted from 16 to 23 May 2009 duringthe beginning of the dry season The savannahlakes were sampled only from the shoreline or theoutflow (Lake Antagnavo) because of the dangerof crocodiles In the other lakes the maximumdepth was measured from an inflatable dinghywith a portable gauge Depth-specific watersamples were collected at the deepest point with aSchindler-Patalas trap and profiles of temperature(thermometer within trap) pH conductivity(Hanna Combo electrode) and oxygen (Hach HQ40d) were measured Integrated qualitative planktonsamples were taken with a 30mm plankton net byretrieving the net from near bottom layers orthrowing out the net from the shore In additionsamples were collected in different littoral habitats(sediment macrophytes) All samples were filteredthrough a 30mm net and preserved in 4formaldehyde Quantitative phytoplankton sampleswere collected from the water surface at the deepestpoint in lakes Mahasarika Malio and Taranta(200mL unfiltered lake water) and at threedifferent depths in lakes Manonja (1 5 9m depth)and Fantany (1 7 15m) and preserved withLugolrsquos solution

In the laboratory Lugol-preserved phytoplanktonsamples were analysed following recommendationsfor counting and biovolume calculations using theUtermoumlhl technique (Rott 1981) Species weredetermined from Lugol and formaldehyde-preservedplankton samples (algae in general E RottZygnematophyceae R Lenzenweger and E Rott)Zooplankton and zoobenthos samples were stainedwith Rose Bengal sorted under a stereo microscopeand determined to genus or species (hydrozoansRD Campbell nematodes W Traunspurgerrotifers R Schabetsberger annelids S Gaviriacladocerans A Kotov copepods F Fiers ostracodsC Meisch mites S Mahunka)

Species assemblages were analysed with non-linearmultidimensional scaling (NMDS Kruskal 1964)using presenceabsence data of all taxa occurringin more than one lake (103 out of 276 taxa mean215 153 SD) and the Kulczynski index (Faithet al 1987 Legendre and Legendre 1998) as ameasure of dissimilarity Using the same index anagglomerative hierarchical clustering of the datawas computed (Kaufman and Rousseeuw 1990)Ordination diagrams and fitted environmentalvectors as well as clustering solutions indicated astrong pattern related to conductivitySubsequently an Analysis of Variance UsingDistance Matrices (Anderson 2001 McArdle andAnderson 2001) was applied to test the nullhypothesis of no relationship between speciesassemblages and conductivity All computationswere coded in R (R Development Core Team2011) using base packages plus the vegan library(Oksanen et al 2011)

RESULTS

Description of lakes

Data on physical conditions and algae are presentedin Figures 3 and 4

Mahasarika (1 Figure 2(a))

The 48m deep crater lake is oversaturated withoxygen down to 4m depth A strong gradient inpH and high total algal biomass result from abloom of Monoraphidium contortum Microcystiswesenbergii Cylindrocystis sp and Cosmarium spIt is said that the lake feeds the water supplyfor the town of Antsiranana (Diego-SuarezA Amba personal communication) although to

Figure 1 Location of the sampling sites in the Amber MountainsNational Park and surroundings (green forested (lakes 1 2 3 7 8)

red savannah (4 5 6))



our knowledge there are no hydrological studiesavailable to confirm this

Malio (2 Figure 2(b))

The highest lake within the National Park wassaturated with oxygen throughout the 4m deepwater column Weak gradients in all parametersindicated frequent mixing of the shallow lakePhytoplankton biomass was dominated byCyanobacteria (Cyanodictyon sp Microcystispulverea Microcystis sp) and Dinophyceae(Peridinium volzii)

Taranta (3 Figure 2(c))

The lake is only 09m deep and well oxygenatedPhytoplankton was dominated by Zygenematophyceae(Cosmarium pygmaeum) Chlorophyceae (Coelastrumsphaericum) and Dinophyceae (Peridiniopsiselpatiewskyi)

Mahery (4 Figure 2(d))

Several small islands indicate a shallow depth of thelake (probablylt 3m) The near-shore water ischaracterized by comparatively higher conductivity

Figure 2 Photographs of the lakes in the Amber Mountains National Park and surroundings



and slightly acidic conditions within a dense belt ofthe non-native water hyacinth Eichhornia crassipes

Antagnavo ely (5 Figure 2(e))

The small crater lake in the south east of LakeAntagnavo is also surrounded by floating carpetsof E crassipes Similar to Lake Mahery surfaceconductivity wasgt 200 mS cm-1 and pH was justabove neutral

Antagnavo (6 Figure 2(f))

The water in the outflow was characterized by highpH and conductivity (Table 1) Crocodiles are fedregularly in ritual ceremonies

Manonja (7 Figure 2(g))

Lake Manonja is the most pristine freshwaterecosystem in the Amber Mountains NationalPark The protected crater lake is 25m deep andexhibited steep physico-chemical gradientsBeyond 15m depth the water was anoxic Aphytoplankton bloom resulted in high pH valuesin shallow water The algal community wasdominated by Cyanobacteria (Anabaena spCyanodictyon sp) So far the lake has not beenstocked with alien fish and is said to contain largenumbers of native eels (Anguilla sp A Ambapersonal communication) Plantations of lsquokhatrsquo(Catha edulis) are only an hourrsquos walk away fromthe lake and numerous felled rosewood trees wereseen in the area

Fantany (8 Figure 2(h))

The deepest lake in the Amber Mountains NationalPark (32m) is situated at the border betweenrainforest and savannah Similar to Lake Manonjasteep physico-chemical gradients were found andthe water was anoxic beyond 15m depthPhytoplankton was dominated by Cyanobacteria(Microcystis pulverea M wesenbergii Cyanothecesp) Bacillariophyceae (Synedra sp) andDinophyceae (Peridiniopsis elpatiewskyi) Accordingto information from local people crocodiles havebeen introduced into the lake but so far no fishhave been stocked The illegal harvest of hardwoodwas witnessed near the lake

Species assemblages

In total 142 algal 1 hydrozoan 23 nematode 56rotifer 12 annelid 15 copepod 16 cladoceran 8ostracod and 3 mite taxa were recordedT

able

1Description

oflakesin

theAmberMou

ntains

Nationa

lPark

Mad

agascar(indicatesna

tive

fish

species)

Lak

eNum

ber

Malagasy

Nam

eForeign

Nam

eSam

pling

Date

Altitude

(m)

Max

Length

(m)

Max

Depth

(m)

Tem

p(C)

Cond

(mScm

-1)

pHOxy

gen

(mgL-1)

Oxy

gen

(sat)

Fish

1Mah

asarika

PetitLac

16052009

1060

280

48

246

28928

824

115

Tila

piarendalli

Cyprinuscarpio

Micropterus

salm

oides

Gam

busiaaffinis

2Malio

Grand

Lac

16052009

1335

360

40

206

31841

797

105

CyprinuscarpioM

icropterus

salm

oides

3Taran

taMau

dit

18052009

1250

720

09

198

56822

809

103

CyprinuscarpioM

icropterus

salm

oides

4Mah

ery

Mah

ery

19052009

364

765

Shallow

261

218

687

--

Tila

piarendalli

Paratila

piasp

5Antagna

voely

Sacreacutesm

all

20052009

348

190

25

259

202

716

--

Tila

piarendalli

Paratila

piasp

Ptychochrom

issp

6Antagna

voSa

creacute

20052009

367

1660

261

231

874

Tila

piarendalli

Paratila

piasp

Ptychochrom

issp

7Man

onja

Texier

22052009

1034

630

25227

61901

872

113

nointrod

uced

fishe

els

8Fan

tany

Fan

tany

23052009

810

780

32254

75906

801

108

nointrod

uced

fishe

els



(Appendix) The highest species richness wasobserved in the belt of water hyacinth of LakeMahery (4) (Figure 5) Crustacean speciesexhibited a lower proportion of cosmopolitan orwidely distributed taxa than the other groups oforganisms Truly endemic morpho-species (3) werepredominantly found within the copepods andthey were restricted to lakes within the parkboundaries (Appendix)

Ordinations and clustering solutions clearlycorroborate differences in species assemblages withregard to conductivity and surrounding vegetation

(Figure 6) Conductivity was significantly relatedto vegetation (one-way ANOVA F16 152 Plt00001) with mean 502 mS cm-1 202 SD forforest and 217 mS cm-1 145 for savannahenvironments Analysis of Variance UsingDistance Matrices confirmed that conductivity isnon-randomly related to species patterns andexplains roughly one-third of the variability in thespecies matrix (R-squared 034 F16 305 P0003 P-values obtained by 999 permutations)The same holds true for surrounding vegetation(2-level factor forest and savannah) when used as

Figure 4 Phytoplankton biomass (mm3L-1) at 05m depth in shallow lakes (top panels) and at three different depths in the deeper lakes (gt 5m bottompanels) within the Amber Mountains National Park

Figure 3 Depth profiles of temperature conductivity pH and oxygen in four lakes of the Amber Mountains National Park



predictor variable (R-squared 033 F16 298P 0016 999 permutations) The effect of introducedfish on species assemblages and richness was alsotested Analysis of Variance was applied using

Distance Matrices and one-way ANOVA oflog-transformed species richness Only spuriousrelationships were found

Ordination of lake communities (Figure 6(a))shows a distinct pattern with regard to conductivityLikewise items display a clear group structurewhen ordination scores are labelled according to thesurrounding environment Independent clusteranalyses (Figure 6(b)) corroborate the validity ofthe ordination configuration In addition clusteringanalyses suggest a hierarchical structure in the dataembedding the three savannah lakes in a group withtwo forest lakes of intermediate (7) and lowconductivity (2)

DISCUSSION

Species communities differed between lakes situatedin forested compared with deforested catchmentsIn turn forest cover was strongly correlated withwater conductivity being four times higher inlowland lakes surrounded by savannah Increasedsediment input through soil erosion probably leadsto changes in algal and micrometazoancommunities (Lott et al 1994 Alin et al 2002)although single sampling cannot provideconclusive evidence Deforestation proceeded fromthe lowlands around Lake Antagnavo (6) towardsthe foothills of the Amber Mountains within thelast millennium (Burney et al 2004) and has nowreached Lake Fantany (8) We hypothesize thatsediment cores would reveal clear changes inspecies communities after the arrival of humansthat are not paralleled in the forest lakes In thisrespect the lakes would be promising targets forpalaeolimnological research

Figure 5 Total number of species grouped at higher taxonomic level(Appendix) The stack bars give the number of species occurring inmore than one lake (solid black) and the fraction of species

encountered only within the named lake (white)

Figure 6 Ordination diagram and cluster dendrogram based on the Kulczynski index computed with presenceabsence data a NMDS Ordination(stress 785) with sample sites connected by a cluster dendrogram The star denotes the root of the dendrogram (see b) The fit of conductivityonto the ordination is shown as a symbolic arrow (R-squared 091 P 0002 p-values based on 999 permutations) b Agglomerative Nesting

Clustering (agglomerative coefficient 015) with tips coded according to the surrounding vegetation



Owing to their sheltered position and nutrient-richvolcanic bedrock all deeper crater lakes withinthe park exhibited strong physico-chemicalgradients anoxic hypolimnia and phytoplanktonblooms The highest epilimnetic algal biomasswas recorded in the lake with four introducedalien fish species This might have resulted fromthe absence of larger crustacean grazers becauseof size-selective fish predation Unfortunatelyquantitative phytoplankton samples from lowlandlakes were lost

No significant differences were detected betweenspecies assemblages of the stocked and the twounstocked lakes The effects of fish were probablymasked by more prominent factors such as forestcover In the highest lake of the National Parkintroduced fish coexisted with the only endemiccalanoid copepod species Tropodiaptomusmadagascariensis This is different from the resultsobtained from a high-altitude lake of similar sizeand depth where native calanoids were eliminatedwithin a decade after stocking (Schabetsbergeret al 2009b see also Knapp et al 2001) Either thelake was stocked fairly recently by carrying fishfrom the lower lakes uphill or the copepod cantolerate size-selective fish predation However westill postulate that palaeolimnological work wouldreveal changes within the species communities afterfish introduction especially as the introducedcyprinids tilapias and mosquitofish prey ondifferent trophic levels from detritus to algae toinsect larvae (Zambrano et al 1999 Fishbase 2012)

Deforestation and introduction of alien speciesdoes not necessarily result in a decrease in speciesrichness For example the large belt of Eichhorniain Lake Mahery (4) probably decreases the pH toslightly acidic conditions through CO2

accumulation (Rai and Munshi 1979) enablingthe development of a rich desmid flora surpassingany other sampling site in its diversity Howeverthe high proportion of taxa that were recordedonly once suggests that the single preliminarysurvey does not warrant a comparison of overallspecies richness between lakes especially as thelargest crater lake could only be sampled in theoutflow

Similar to other isolated crater lakes onMadagascar islands of Oceania and in WestAfrica (Schabetsberger et al 2004 2009a c) alarge proportion of species were cosmopolitansor widely distributed taxa Only within thecrustaceans did tropicopolitan and afrotropicalfaunal elements predominate The Mozambique

Channel may not be a barrier for the transport oftheir propagules with wind rain and water birdsTruly endemic species were predominantly foundwithin the copepods which seem to have limiteddispersal abilities (Schabetsberger et al 2009c)

On the other hand there is growing evidencethat many freshwater algal and micrometazoanmorpho-species are actually complexes of crypticspecies with more restricted distributions (Goacutemezet al 2002 Fawley et al 2004 Belyaeva andTaylor 2009) In addition some species that couldonly be determined to genus level may be newendemic taxa Hence remote lakes on islands maystill harbour comparatively higher proportions ofgenetically isolated lineages of microorganisms(Schroumlder and Walsh 2007) With several species ofhigher plants and animals on the brink of extinctionin Madagascar conserving potentially undescribedplanktonic and benthic organisms has low priorityAt present the lakes attract internationalconservation efforts only if lsquoflagshiprsquo bird speciesoccur (eg the Madagascar pochard Aythyainnotata (TWSG News 2006)) With the currentstate of knowledge it is impossible to understanddistribution patterns and the full degree ofendemicity in Malagasy freshwater organisms

Conserving the lakes and their catchmentsfrom further deterioration is difficult The AmberMountains and their lakes are protected as aninternationally recognized National Park which isthe highest status of protection in MadagascarHowever the raiding of the forests is highlyorganized and well funded as 1m3 of rosewoodyields US$ 3000 (Wikipedia 2012) Encampments ofloggers may invite the introduction of alien fish intothe last unstocked lake The eradication of alien fishpopulations by intensive gill netting and seining(Knapp and Matthews 1998) might helptemporarily to restore more natural conditions in theshallow already stocked lakes and could be achievedwith comparatively little funding However as longas some precious timber can leave the country theremaining forests will be steadily depleted and theimpact on lake ecosystems will increase

The protection of National Parks from illegaltimber harvest is difficult and requires frequentcontrols of armed rangers which is unrealistic underthe current political situation Owing to internationalpressure timber exports were banned in 2010(Bohannon 2010) but on 18 January 2012Madagascarrsquos Minister of the Environment hasre-authorized the export of lsquoall categories of naturalforest-sourced primary productsrsquo (Mongabay 2012)



Only international loans financing conservationefforts in Madagascar coupled with political pressuremay ease the current situation although it is likelyonly to slow down the depletion until the trees finallybecome too scarce to be harvested economicallySubsequently slash-and-burn agriculture treecutting honey extraction and bushmeat huntingwill commence within the degraded catchments Bythen the Amber Mountains lake communities willresemble those from lowland areas

ACKNOWLEDGEMENTS

We thankAAmba CHolmes A Randimbison andRS Harison for support of the field operation CJersabek and two anonymous reviewers providednumerous corrections The work was conductedunder the permit lsquoAutorisation de recherche en eaudouce 02ARED09rsquo issued by the Ministegravere delrsquoAgriculture de lrsquoEacutelevage et de la Pecircche

REFERENCES

Alin RS OrsquoReilly M Cohen AS Dettman DL Palacios-FestMR McKee BA 2002 Effects of land use change onaquatic biodiversity a view from the paleorecord of LakeTanganyika East Africa Geology 30 1143ndash1146

Anderson M 2001 A new method for non-parametricmultivariate analysis of variance Austral Ecology 26 32ndash46

Belyaeva M Taylor DJ 2009 Cryptic species within theChydorus sphaericus species complex (CrustaceaCladocera)revealed by molecular markers and sexual stagemorphology Molecular Phylogenetics and Evolution 50534ndash546

Benstead JP DeRham PH Gattolliat J-L Gibon F-MLoiselle PV Sartori M Sparks JS Stiassny MLJ 2003Conserving Madagascarrsquos freshwater biodiversityBioScience 53 1101ndash1111

Berzins B 1960 Neue Rotatorienarten aus MadagaskarMeacutemoires de la Institut Scientifique de Madagascar 14 1ndash6

Berzins B 1973 Zwei neue Euchlanis ndashArten RotatoriaZoologischer Anzeiger 190 125ndash127

Berzins B 1982 Zur Kenntnis der RotatorienfaunaMadagaskars AV centralen i Lund Lund Sweden

Bohannon J 2010 Madagascarrsquos forests get a reprieve ndash butfor how long Science 328 23ndash25

Bourrelly P Leboime R 1946 Notes sur quelques alguesdrsquoeau douce de Madagascar Biologisch Jaarbock Dodonea13 75ndash111

Bourrelly P Manguin E 1949 Contribution agrave lrsquoeacutetude de laflore algale drsquoeau douce de Madagascar Le lac deTzimbazaza Meacutemoires de la Institut Scientifique deMadagascar 2 161ndash190

Bourrelly P Couteacute A 1991 Desmidieacutees de MadagascarBibliotheca Phycologica 86 Gebruumlder Borntraeger Berlin-Stuttgart

Brehm V 1930 Notizen zur Cladocerenfauna MadagaskarsArchiv fuumlr Hydrobiologie 21 679ndash686

BrehmV 1948 Nuevos copepodos deMadagascarPublicacionesInstituto de Biologia applicada Barcelona 5 77ndash84

Brehm V 1951 Pseudodiaptomus pauliani Der erste Vertreterder Pseudodiaptomiden in der madegassichen FaunaMeacutemoires de la Institut Scientifique deMadagascar 6 419ndash425

Brehm V 1952a Cladoceren und calanoide Copepoden vonMadagaskar Meacutemoires de la Institut Scientifique deMadagascar 7 37ndash46

Brehm V 1952b Anadiaptomus madagascariensis Rylov(Copepoda Diaptomidae) Le Naturaliste Malgache 3159ndash162

Brehm V 1952c Anadiaptomus poseidon nov gen nov specaus Madagaskar Anzeiger der oumlsterreichischen Akademieder Wissenschaften 89 23ndash27

Brehm V 1953 Cladocegraveres et copeacutepodes calanoides deMadagascar Le Naturaliste Malgache 5 151ndash152

Brehm V 1954 Pseudodiaptomus batillipes spec nov einzweiter Pseudodiaptomus aus Madagaskar Sitzungsberichteder oumlsterreichischen Akademie der WissenschaftenMathematisch ndash Naturwissenschaftliche Klasse 1 603ndash607

Brehm V 1960 Ergebnisse der oumlsterreichischen Madagaskar-Expedition 1958 1 Beitrag zur Kenntnis der Crustaceamadegassischer Stillgewaumlsser Meacutemoires de la InstitutScientifique de Madagascar 14 39ndash58

Burney DA Burney LP Godfrey LR Jungers WL GoodmannSM Wright HT Jull AJT 2004 A chronology for lateprehistoric Madagascar Journal of Human Evolution 4725ndash63

Canonico GC Arthington A McCrary JK Thieme ML 2005The effects of introduced tilapias on native biodiversityAquatic Conservation Marine and Freshwater Ecosystems15 463ndash483

Chappuis PA 1952 Copeacutepodes harpacticoides psammiques deMadagascar Meacutemoires de la Institut Scientifique deMadagascar 7 145ndash160

Chappuis PA 1954 Recherches sur la faune interstitielle dessediment marins et drsquoeau douce a Madagascar IVCopeacutepodes harpacticoides psammiques de MadagascarMeacutemoires de la Institut Scientifique de Madagascar 9 45ndash73

Chappuis PA 1956 Presence a Madagascar du genreEchinocamptus E pauliani nsp Meacutemoires de la InstitutScientifique de Madagascar 10 35ndash42

Coesel PFM 2002 Taxonomic and biogeographical notes onMalagasy desmids (Chlorophyta Desmidiaceae) NordicJournal of Botany 22 239ndash256

Cucciniello C Melluso L Morra V Storey M Rocco IFranciosi L Grifa C Petrone CM Vincent M 2011 New40Ar-39Ar ages and petrogenesis of the Massif drsquoAmbrevolcano northern Madagascar Geological Society ofAmerica Special Papers 478 257ndash281

DeGuerne J Richard J 1893 Canthocamptus grandidieriAlona cambouei nouveaux entomostraces drsquoeau douce deMadagascar Meacutemoires de la Societeacute zoologique de France96 234ndash244

Dussart BH 1982 Crustaceacutes Copegravepodes des eaux inteacuterieuresFaune de Madagascar 58 ORSTOM CNRS Paris

Faith FP Minchin PR Belbin L 1987 Compositionaldissimilarity as a robust measure of ecological distanceVegetatio 69 57ndash68

Fawley MW Fawley KP Buchheim MA 2004 Moleculardiversity among communities of freshwater microchlorophytesMicrobial Ecology 48 489ndash499

Fiers F 2002 The genus Haplocyclops Kiefer 1952(Copepoda Cyclopoida Cyclopidae) redescription of thetype-species H gudrunae and its congeners Hydrobiologia474 155ndash169

Fishbase 2012 httpwwwfishbaseorg (30 March 2012)Fritsch FE 1914 Contribution to our knowledge of thefreshwater algae of Africa I Some freshwater algae fromMadagascar Annales de Biologie Lacustre 7 40ndash59



Ganzhorn JU Lowry PP Schatz GE Sommer S 2008 Thebiodiversity of Madagascar one of the worldrsquos hottesthotspots on its way out Oryx 35 346ndash348

Goacutemez A Serra M Carvalho GR Lunt DH 2002 Speciationin ancient cryptic species complexes evidence from themolecular phylogeny of Brachionus plicatilis (Rotifera)Evolution 56 1431ndash1444

Groombridge B Jenkins M 1998 Freshwater Biodiversity APreliminary Global Asessment World ConservationMonitoring Centre (WCMC) Biodiversity Series no 8World Conservation Press WCMC Cambridge

Hansen P 1996 Silica-scaledChrysophyceae and SynurophyceaefromMadagascar Archiv fuumlr Protistenkunde 147 145ndash172

Harper GJ Steininger MC Tucker CJ Juhn D Hawkins F2007 Fifty years of deforestation and forest fragmentationin Madagascar Environmental Conservation 34 325ndash333

Kaufman L Rousseeuw PJ 1990 Finding Groups in Data anIntroduction to Cluster Analysis Wiley New York

Kiefer F 1930 Zur Kenntnis freilebender CopepodenMadagaskars Zoologischer Anzeiger 87 42ndash46

Kiefer F 1952Haplocyclops gudrunae ngen et nsp ein neuerRuderfuszligkrebs (Crustacea Copepoda) aus MadagaskarZoologischer Anzeiger 149 240ndash243

Kiefer F 1954 Neue Cyclopoida Gnathostoma (CrustaceaCopepoda) aus Madagaskar I Cyclopininae andHalicyclopinae Zoologischer Anzeiger 153 308ndash313

Kiefer F 1955 Neue Cyclopoida Gnathostoma (CrustaceaCopepoda) aus Madagaskar II Cyclopinae ZoologischerAnzeiger 154 222ndash232

Knapp RA Matthews KR 1998 Eradication of nonnative fishby gill-netting from a small mountain lake in CaliforniaRestoration Ecology 6 207ndash213

Knapp RA Matthews KR Sarnelle O 2001 Resistence andresilience of Alpine lake fauna to fish introductionsEcological Monographs 71 401ndash421

Kruskal JB 1964 Nonmetric multidimensional scaling anumerical method Psychometrika 29 115ndash129

Legendre P Legendre L 1998 Numerical Ecology 2nd ednElsevier Amsterdam

Lindberg K 1951a Cyclopides de Madagascar I Meacutemoires dela Institut Scientifique de Madagascar 5 187ndash195

Lindberg K 1951b Cyclopides de Madagascar II Meacutemoiresde la Institut Scientifique de Madagascar 6 427ndash437

Lindberg K 1952 Cyclopides de Madagascar III Meacutemoiresde la Institut Scientifique de Madagascar 7 53ndash67

Lindberg K 1953 Cyclopides de Madagascar IV Meacutemoires dela Institut Scientifique de Madagascar 8 11ndash17

Lott A-M Siver PA Mariscano LJ Kodoma KP Moeller RE1994 The paleolimnology of a small waterbody in thePocono Mountains of Pennsylvania USA reconstructing19thndash20th century specific conductivity trends in relation tochanging land use Journal of Paleolimnology 12 75ndash86

Manguin E 1941 Contribution agrave la connaisance desDiatomeacutees drsquoeau douce de Madagascar Revue Algologique12 153ndash157

McArdle B Anderson M 2001 Fitting multivariate models tocommunity data a comment on distance-based redundancyanalysis Ecology 82 290ndash297

Metzeltin D LangendashBertalot H 2002 Diatoms from the ldquoIslandContinentrdquo Madagascar Iconographia Diatomologica 11Koeltz Scientific Books Koumlnigstein

Mongabay 2011 httprainforestsmongabaycom20madagascarhtm (2 November 2011)

Mongabay 2012 httpnewsmongabaycom20120229-rosewood_ban_liftedhtml (30 March 2012)

Myers N Mittermeier RA Mittermeier CG Da FonsecaGAB Kent J 2000 Biodiversity hotspots for conservationpriorities Nature 403 853ndash858

Oksanen J Guillaume Blanchet F Kindt R Legendre PMinchin PR OrsquoHara RB Simpson GL Solymos PStevens MHH Wagner H 2011 Vegan CommunityEcology Package R version 20-0 httpveganr-forger-projectorg

Rai DN Munshi JD 1979 The influence of thick floatingvegetation (Water hyacinth Eichhornia crassipes) on thephysico-chemical environment of a freshwater wetlandHydrobiologia 62 65ndash69

Raxworthy CJ Pearson RG Rabibisoa N RakotodrazafyAM Ramanamanjato J-P Raselimanana AP Wu SNussbaum RA Stone DA 2008 Extinction vulnerability oftropical montane endemism from warming and upslopedisplacement a preliminary appraisal for the highest massifin Madagascar Global Change Biology 14 1703ndash1720

R Development Core Team 2011 R A Language andEnvironment for Statistical Computing R Foundation forStatistical Computing Vienna Austria

Reinthal PN Stiassny MLJ 1991 The freshwater fishes ofMadagascar a study of an endangered fauna withrecommendations for a conservation strategy ConservationBiology 5 231ndash243

Rott E 1981 Some results from phytoplankton countingintercalibrations Schweizerische Zeitschrift fuumlr Hydrologie43 34ndash63

Sala OE Chapin FS Armesto JJ Berlow E Bloomfield JDirzo R Huber-Sanwald E Huenneke LF Jackson RBKinzig A et al 2000 Biodiversity ndash global biodiversityscenarios for the year 2100 Science 287 1770ndash1774

Schabetsberger R Drozdowski G Drozdowski I Jersabek CDRott E 2004 Limnological aspects of two tropical craterlakes (Lago Biao and Lago Loreto) on the island of Bioko(Equatorial Guinea) Hydrobiologia 524 79ndash90

Schabetsberger R Rott E Friedl G Drozdowski GRazafinadranaivo E Holmes C 2009a First limnologicalcharacterization of the tropical crater lake Amparihibein the Makira Protected Area Madagascar Eco-Mont 143ndash52

Schabetsberger R Luger M Drozdowski G Jagsch A 2009bOnly the small survive ndash monitoring long-term changes inthe zooplankton community of an Alpine lake after fishintroduction Biological Invasions 11 1335ndash1345

Schabetsberger R Drozdowski G Rott E Lenzenweger RJersabek CD Fiers F Traunspurger W Reiff N Stoch FKotov AA et al 2009c Losing the bounty Investigatingspecies richness in isolated freshwater ecosystems ofOceania Pacific Science 63 153ndash181

Schroumlder T Walsh EJ 2007 Cryptic speciation in thecosmopolitan Epiphanes senta complex (MonogonontaRotifera) with the description of new species Hydrobiologia593 129ndash140

Segers H 1992 Taxonomy and zoogeography of the rotiferfauna of Madagascar and the Comoros Journal of AfricanZoology 106 351ndash361

Schuurman D Lowry PP 2009 The Madagascar rosewoodmassacre Madagascar Conservation and Development 498ndash102

Sparks JS Stiassny MLJ 2003 Introduction to the freshwaterfishes In The Natural History of Madagascar GoodmanSM Benstead JP (eds) University of Chicago PressChicago 849ndash863

Spaulding SA Kociolek JP 1998a New Gomphonema(Bacillariophyceae) species from Madagascar Proceedingsof the California Academy of Sciences 50 361ndash379

Spaulding SA Kociolek JP 1998b The diatom genusOrthoseira ultrastructure and morphological variation intwo species from Madagascar with comments onnomenclature in the genus Diatom Research 13 133ndash147



TWSG News 2006 Madagascar Pochard rediscovered TheBulletin of the Threatened Waterfowl Specialist Group 15 8

Vences M Wollenberg KC Vietes DR Lees DC 2009Madagascar as a model region of species diversificationTrends in Ecology amp Evolution 24 456ndash465

Vieites DR Wollenberg KC Andreone F Koehler J Glaw FVences M 2009 Vast underestimation of Madagascarrsquosbiodiversity evidenced by an integrative amphibian inventoryProceedings of the National Academy of Sciences USA 1068267ndash8272

West W West GS 1895 A contribution to our knowledgeof the freshwater algae of Madagascar Transactionsof the Linnean Society London Botany 5 41ndash90plates 5ndash9

Wikipedia 2012 httpenwikipediaorgwikiIllegal_logging_in_Madagascar (2 July 2012)

Zambrano L Perrow MR Maciacuteas-Garciacutea C Hidalgo V 1999Impact of introduced carp (Cyprinus carpio) in subtropicalshallow ponds in Central Mexico Journal of AquaticEcosystems Stress and Recovery 6 281ndash288

APPENDIX

Freshwater algae Hydrozoa Nematoda Rotifera Annelida Copepoda Cladocera Ostracoda and Acari found in theeight lakes of the Amber Mountains National Park (1ndash8 Table 1) and surroundings and their known distribution(afrotrop = afrotropical austral = Australian circumtrop = circumtropical cosm = cosmopolitan holarct = holarcticnearct = nearctic neotrop = neotropical orient = oriental palaearct = palaearctic tropicopol = tropicopolitan)Circumtropical distribution predominantly between the Tropics of Cancer (2327rsquo N) and Capricorn (23270 S)Tropicopolitan distribution throughout the tropical and subtropical zone (up to app 34 N and S) but at higherlatitudes if local temperature regimes permit

Group Species NamesLake

Number Biogeography

Cyanophyceae Anabaena sp 127Cyanophyceae Aphanocapsa delicatissima W West amp GS West 1912 13 cosmCyanophyceae Chroococcus limneticus Lemmermann 1898 3 cosmCyanophyceae Chroococcus sp 1Cyanophyceae Cyanodictyon sp 27Cyanophyceae Cyanothece sp 28Cyanophyceae Cylindrospermopsis africana Komaacuterek amp Kling 1991 1 afrotropCyanophyceae Geitlerinema splendidum (Greville) Anagnostidis 1989 4 cosmCyanophyceae Merismopedia tenuissima Lemmermann 1898 1 cosmCyanophyceae Microcystis aeruginosa (Kuumltzing) Kuumltzing 1846 128 cosmCyanophyceae Microcystis comperei Komaacuterek 1984 1Cyanophyceae Microcystis protocystis Crow 1932 8 circumtropCyanophyceae Microcystis pulverea (Wood) Forti 1907 28 cosmCyanophyceae Microcystis wesenbergii (Komaacuterek) Komaacuterek 1968 138 cosmCyanophyceae Microcystis sp 1Cyanophyceae Oscillatoria sp 3Cyanophyceae Planktolyngbya limnetica (Lemmermann)

J Komaacuterkovaacute-Legnerovaacute amp G Cronberg 19921238 cosm

Cyanophyceae Planktolyngbya undulata Komaacuterek amp Kling 1991 3 afrotropCyanophyceae Radiocystis sp 2Cryptophyceae Chroomonas acuta Utermoumlhl 1925 2 cosmCryptophyceae Cryptomonas tetrapyrenoidosa Skuja 1948 2 cosmCryptophyceae Cryptomonas sp 1Dinophyceae Gymnodinium sp 5Dinophyceae Peridiniopsis elpatiewskyi (Ostenfeld) Bourrelly 1968 138 cosmDinophyceae Peridinium cf pusillum (Peacutenard) Lemmermann 1901 8 cosmDinophyceae Peridinium volzii var cinctiforme MLefegravevre 1932 27 endemicDinophyceae Peridinium sp 1 238Dinophyceae Peridinium sp 2 18Dinophyceae Wolozynskia sp 5Chrysophyceae Mallomonas sp 5Chrysophyceae Spiniferomonas sp 5Bacillariophyceae Aulacoseira ambigua (Grunow) Simonsen 1979 128 cosmBacillariophyceae Aulacoseira granulata (Ehrenberg) Simonsen 1979 137 cosmBacillariophyceae Cyclotella meneghiniana Kuumltzing 1844 2 cosmBacillariophyceae Cyclotella sp 1Bacillariophyceae Surirella sp 6Bacillariophyceae Synedra sp 18Chlorophyceae Ankistrodesmus bernardii Komaacuterek 1983 4 circumtropChlorophyceae Asterococcus sp 4Chlorophyceae Botryococcus sp 237Chlorophyceae Chlamydomonas sp 1Chlorophyceae Chlorolobion (Ankistrodesmus) braunii (Naumlgeli) Komaacuterek 1979 4 cosm

(Continues)



Table 1 Continued

Group Species NamesLakeNumber Biogeography

Chlorophyceae Chlorotetraedron incus (Teiling) Komaacuterek amp Kovaacutecik 1985 13 cosmChlorophyceae Coelastrum indicum WB Turner 1892 4 tropicopolChlorophyceae Coelastrum pulchrum Schmidle 1892 1Chlorophyceae Coelastrum proboscideum Bohlin in Wittrock amp Nordstedt 1896 1 cosmChlorophyceae Coelastrum reticulatum (PA Dangeard) Senn 1899 4 cosmChlorophyceae Coelastrum reticulatum var cubanum Komaacuterek 1975 38 circumtropChlorophyceae Coelastrum sphaericum Naumlgeli 1849 3 cosmChlorophyceae Coenococcus sp 478Chlorophyceae Crucigenia tetrapedia W West amp GS West 1902 2 cosmChlorophyceae Dictyosphaerium tetrachotomum var fallax Komaacuterek 1983 3 circumtropChlorophyceae Dictyosphaerium sp 1 48Chlorophyceae Dictyosphaerium sp 2 1Chlorophyceae Golenkinia sp 1Chlorophyceae Kirchneriella dianae (Bohlin) Comas Gonzalez 1980 134 tropicopolChlorophyceae Lagerheimia longiseta (Lemmermann) Printz 1914 1 cosmChlorophyceae Lauterborniella sp 1Chlorophyceae Monoraphidium contortum (Thuret)

Komagraverkovaacute-Legnerovaacute 1969128 cosm

Chlorophyceae Oocystis borgei JSnow 1903 2 cosmChlorophyceae Oocystis sp 48Chlorophyceae Pandorina morum (OF Muumlller) Bory de Saint-Vincent 1824 4 cosmChlorophyceae Pediastrum duplex Meyen 1829 125 cosmChlorophyceae Pediastrum duplex var echinatum CC Jao 1947 1 cosmChlorophyceae Pediastrum simplex Meyen 1829 3 cosmChlorophyceae Pediastrum tetras (Ehrenberg) Ralfs 1844 1248 cosmChlorophyceae Pseudodictyosphaerium elegans (Bachmann) Hindaacutek 1988 45 cosmChlorophyceae Scenedesmus acuminatus (Lagerheim) Chodat 1902 12 cosmChlorophyceae Scenedesmus armatus (R Chodat) R Chodat 1913 1Chlorophyceae Scenedesmus brasiliensis Bohlin 1897 2 cosmChlorophyceae Scenedesmus opoliensis PG Richter 1897 23 cosmChlorophyceae Scenedesmus tropicus WB Crow 1923 1 tropicopolChlorophyceae Scenedesmus sp 1 38Chlorophyceae Scenedesmus sp 2 1Chlorophyceae Scourfieldia sp 18Chlorophyceae Sorastrum sp 4Chlorophyceae Tetraedron minimum (ABraun) Hansgirg 1888 18 cosmChlorophyceae Tetrastrum staurogeniaeforme (Schroumlder) Lemmermann 1900 17 cosmZygnematophyceae Bambusina brebissonii Kuumltzing 1945 4 cosmZygnematophyceae Closterium dianae var arcuatum (Breacutebisson ex Ralfs)

Rabenhorst 18682 cosm

Zygnematophyceae Closterium ehrenbergii Meneghini ex Ralfs 1848 4 cosmZygnematophyceae Closterium lineatum Ehrenberg ex Ralfs 1848 4 cosmZygnematophyceae Closterium parvulum Naumlgeli 1849 4 cosmZygnematophyceae Closterium ralfsii Breacutebisson ex Breacutebisson in Ralfs 1848 4 cosmZygnematophyceae Closterium setaceum Ehrenberg ex Ralfs 1848 4 cosmZygnematophyceae Cosmarium binum Nordstedt in Wittrock amp Nordstedt 1880 4 cosmZygnematophyceae Cosmarium contractum var minutum (Delponte) Coesel 1989 1 4 8 cosmZygnematophyceae Cosmarium depressum (Naumlgeli) PLundell 1871 1 cosmZygnematophyceae Cosmarium nudum (WB Turner) Gutwinski 1902 4 tropicopolZygnematophyceae Cosmarium pygmaeum WArcher 1864 38 cosmZygnematophyceae Cosmarium quadrum var sublatum (Nordstedt)

W West amp GS West 19124 cosm

Zygnematophyceae Cosmarium spyridion W West amp GS West 1895 14 endemicZygnematophyceae Cosmarium sp 2 2Zygnematophyceae Cosmarium sp 3 4 endemicZygnematophyceae Cylindrocystis sp 12Zygnematophyceae Euastrum didelta Ralfs ex Ralfs 1848 4 cosmZygnematophyceae Euastrum elegans var madagascariense W West amp GS West 1895 4 cosmZygnematophyceae Euastrum humerosum Ralfs 1848 4 cosmZygnematophyceae Euastrum praemorsum (Nordstedt) Schmidle 1898 4 cosmZygnematophyceae Euastrum sp 145Zygnematophyceae Gonatozygon sp 4Zygnematophyceae Micrasterias anomale Turner 1892 4 tropicopolZygnematophyceae Micrasterias crux-melitensis (Ehrenberg) Hassall ex Ralfs 1848 4 cosmZygnematophyceae Micrasterias decemdentata (Naumlgeli) WArcher 1861 4 cosmZygnematophyceae Micrasterias truncata var africana FE Fritsch amp MF Rich 1924 4 tropicopolZygnematophyceae Mougeotia sp 4Zygnematophyceae Pleurotaenium ehrenbergii (Breacutebisson ex Ralfs) Delponte 1878 4 cosm

(Continues)

(Continued)



Table 1 Continued


Zygnematophyceae Pleurotaenium ovatum Nordstedt 1877 4 tropicopolZygnematophyceae Pleurotaenium trabecula var rectum (Delponte) W West amp GS

West 19044 cosm

Zygnematophyceae Staurastrum brachiatum Ralfs ex Ralfs 1848 8 cosmZygnematophyceae Staurastrum ceylanicum W West amp GS West 1902 2 orientZygnematophyceae Staurastrum ellipticum West 1892 8 cosmZygnematophyceae Staurastrum gracile var coronulatum Boldt 1885 8 cosmZygnematophyceae Staurastrum inflexum Breacutebisson 1856 4 cosmZygnematophyceae Staurastrum johnsonii W West amp GS West 1896 1 cosmZygnematophyceae Staurastrum leptocladum Nordstedt 1869 4 tropicopolZygnematophyceae Staurastrum muticum Breacutebisson ex Ralfs 1848 27 cosmZygnematophyceae Staurastrum orbiculare var ralfsii W West amp GS West 1912 4 cosmZygnematophyceae Staurastrum sebaldi var ornatum Nordstedt 1873 8 cosmZygnematophyceae Staurastrum sp 1 18Zygnematophyceae Staurastrum sp 2 17Zygnematophyceae Staurastrum sp 3 2Zygnematophyceae Staurastrum sp 4 17Zygnematophyceae Staurastrum sp 5 1Zygnematophyceae Staurastrum subavicula (West) W West amp GS West 1894 4 cosmZygnematophyceae Staurastrum tetracerum Ralfs ex Ralfs 1848 138 cosmZygnematophyceae Staurodesmus dejectus (Breacutebisson) Teiling 1967 1 cosmZygnematophyceae Staurodesmus sp 1 3Zygnematophyceae Staurodesmus sp 2 1Euglenophyceae Euglena sp 1 5Euglenophyceae Euglena sp 2 45Euglenophyceae Euglena acus (OFMuumlller) Ehrenberg 1830 5Euglenophyceae Phacus curvicauda Svirenko 1915 34 cosmEuglenophyceae Phacus sp 5Euglenophyceae Strombomonas sp 4Euglenophyceae Trachelomonas horrida Palmer 1905 4 cosmEuglenophyceae Trachelomonas volvocina Ehrenberg 1833 12 cosmEuglenophyceae Trachelomonas sp 34578Xanthophyceae Goniochloris fallax Fott 1960 5 cosmXanthophyceae Trochiscia sp 3Hydrozoa Hydra viridissima Pallas 1766 4 cosmNematoda Achromadora micoletzkyi Stefanski 1915 7 cosmNematoda Crocodorylaimus flavomaculatus (Linstow 1876) 234578 cosmNematoda Cryptonchus sp 2Nematoda Dorylaimus cf stagnalis Dujardin 1845 7 cosmNematoda Eumonhystera cf gerlachi (Meyl 1954) 5 EuropeNematoda Ironus ignavus Bastian 1865 7 cosmNematoda Ironus tenuicaudatus De Man 1876 278 palaearct afrotrop orientNematoda Mesodoryilaimus cf subtiloides (Paetzold 1958) 257 EuropeNematoda Mesodorylaimus sp 1 2Nematoda Mesodorylaimus sp 2 2Nematoda Monhystera cf paludicola De Man 1881 8 cosmNematoda Monhystrella cf paramacrura (Meyl 1953) 7 palaearct afrotrop orientNematoda Mononchus cf aquaticus Coetzee 1968 8 cosmNematoda Mylonchulus lacustris (Cobb 1915) 237 palaearct afrotrop orient nearct

neotropNematoda Neoactinolaimus duplicidentatus (Andraacutessy 1968) 237 palaearct afrotropNematoda Neotobrilus longus (Leidy 1852) 7 palaearct afrotrop orient

nearct neotropNematoda Oncholaimus cf oxyuris Ditlevsen 1911 7 brackish waterNematoda Plectus cf pusillus Cobb 1893 7 palaearct afrotrop austral

nearct neotropNematoda Prodorylaimus sp 7Nematoda Rhabdolaimus aquaticus De Man 1880 2 cosmNematoda Rhabdolaimus terrestris De Man 1880 5 cosmNematoda Species 1 5Nematoda Tobrilus helveticus (Hofmaumlnner 1914) 2457 palaearctRotifera Anuraeopsis fissa Gosse 1851 1568 cosmRotifera Anuraeopsis navicula Rousselet 1911 238 cosmRotifera Asplanchna cf brightwellii Gosse 1850 3 cosmRotifera Brachionus angularis Gosse 1851 138 cosmRotifera Brachionus falcatus Zacharias 1898 125678 cosmRotifera Brachionus quadridentatus Hermann 1783 378 cosmRotifera Cephalodella sp 8

(Continues)

(Continued)



Table 1 Continued


Rotifera Colurella sp 7Rotifera Conochilus sp 7Rotifera Dicranophorus tegillus Harring amp Myers 1928 4 nearc orientRotifera Epiphanes cf clavulata (Ehrenberg 1831) 5 cosmRotifera Euchlanis sp 4Rotifera Filinia opoliensis (Zacharias 1898) 78 cosmRotifera Lepadella (H) ehrenbergi (Perty 1850) 7 cosmRotifera Hexarthra intermedia Wiszniewski 1929 1578 cosmRotifera Keratella procurva (Thorpe 1891) 67 cosm (excl nearct)Rotifera Keratella valga (Ehrenberg 1834) 28 cosmRotifera Lecane curvicornis (Murray 1913) 45 cosmRotifera Lecane bulla (Gosse 1851) 4578 cosmRotifera Lecane aculeate (Jakubski 1912) 3456 cosmRotifera Lecane closterocerca (Schmarda 1859) 57 cosmRotifera Lecane furcata (Murray 1913) 478 cosmRotifera Lecane hamata (Stokes 1896) 4567 cosmRotifera Lecane hornemanni (Ehrenberg 1834) 23 cosmRotifera Lecane inermis (Bryce 1892) 3457 cosmRotifera Lecane inopinata Harring amp Myers 1926 7 cosmRotifera Lecane luna (Muumlller 1776) 48 cosmRotifera Lecane lunaris (Ehrenberg 1832) 2 cosmRotifera Lecane nana (Murray 1913) 4 cosmRotifera Lecane pyriformis (Daday 1905) 467 cosmRotifera Lecane quadridentata Ehrenberg 1830 4 cosmRotifera Lecane cf ruttneri Hauer 1938 6 afrotr orient neotrRotifera Lecane signifera (Voigt 1902) 4 cosmRotifera Lecane thienemanni (Hauer 1938) 6 afrotrop orient austr neotrRotifera Lecane unquitata (Fadeew 1925) 47 eastern hemisphereRotifera Lecane spp 4Rotifera Lepadella patella (Muumlller 1773) 247 cosmRotifera Lepadella sp 5Rotifera Lepadella quinquecostata (Lucks 1912) 4 cosmRotifera Macrochaetus collinsii (Gosse 1867) 4 cosmRotifera Mytilina ventralis (Ehrenberg 1832) 47 cosmRotifera Mytilina sp 4Rotifera Platyias leloupi Gillard 1957 45 pres cosm (excl palaearct)Rotifera Polyarthra indica Segers amp Babu 1999 56 afrotrop orient pacificRotifera Polyarthra vulgaris Carlin 1943 13 cosmRotifera Polyarthra sp 45Rotifera Scaridium sp 4Rotifera Testudinella incisa (Ternetz 1892) 38 cosmRotifera Testudinella patina (Hermann 1783) 458 cosmRotifera Trichocerca cf pusilla (Jennings 1903) 13 cosmRotifera Trichocerca chattoni (De Beauchamp 1907) 1 cosmRotifera Trichocerca cf rattus (Muumlller 1776) 56 cosmRotifera Trichocerca similis (Wierzejski 1893) 1 cosmRotifera Trichocerca tigris (Muumlller 1786) 4 cosmRotifera Trichocerca sp 258Rotifera Trichotria tetractis (Ehrenberg 1830) 2 cosmOligochaeta Dero (Aulophorus) hymanae Naidu 1962 3 Southern IndiaOligochaeta Chaetogaster crystallinus Vejdovsky 1883 7 palaearct afrotrop orient nearctOligochaeta Nais africana Brinkhurst 1966 78 afrotropOligochaeta Nais pseudoobtusa Piguet 1906 3 palaearct afrotrop orient

nearct neotropOligochaeta Nais simplex Piguet 1906 37 palaearct afrotrop orient nearctOligochaeta Nais sp 1 3 endemicOligochaeta Nais sp 2 3Oligochaeta Nais sp 3 3Oligochaeta Pristina aequiseta Bourne 1891 3 cosmOligochaeta Pristina proboscidea Beddard 1896 7 palaearct orient austral neotropOligochaeta Vejdovskyella comata (Vejdovsky 1883) 3 palaearct afrotrop orient nearctHirudinea Alboglossiphonia heteroclita (Linnaeus 1761) 7 palaearct nearctCopepoda Elaphoidella bidens (Schmeil 1894) 38 cosmCopepoda Elaphoidella grandidieri Guerne amp Richard 1893 3 afrotrop orient pacific neotrCopepoda Elaphoidella sp 48Copepoda Epactophanes richardi Mrazek 1893 7 cosmCopepoda Cryptocyclops linjanticus (Kiefer 1928) 45 afrotrop orient holarct

(aralo-caspian and mediterranean)

(Continues)

(Continued)



Table 1 Continued


Copepoda Ectocyclops hirsutus (Kiefer 1930) 4 afrotropCopepoda Mesocyclops insulensis Dussart 1982 8 endemicCopepoda Mesocyclops pilosus (Kiefer1930) 12378 endemicCopepoda Microcyclops cf varicans (Sars 1863) 3457 cosmCopepoda Paracyclops sp 78 Genus cosmopolitanCopepoda Thermocyclops cf crassus (Fischer 1853) 5 cosmCopepoda Thermocyclops decipiens (Kiefer 1929) 14 cosmCopepoda Thermocyclops neglectus (Sars 1909) 6 afrotropCopepoda Tropocyclops cf tenellus (Sars 1909) sensu Lindberg (1952) 8 afrotrop endemicCopepoda Tropodiaptomus (Anadiaptomus) madagascariensis poseidon

Brehm 195227 endemic

Cladocera Alona cambouei Guerne amp Richard 1893 47 circumtrop eastern hemisphereCladocera Alona guttata Sars 1862 8 cosm species groupCladocera Anthalona harti harti Van Damme Sinev amp Dumont 2011 47 afrotropCladocera Ceriodaphnia cornuta Sars 1885 24 tropicopolCladocera Ceriodaphnia laticaudata PE Muumlller 1867 4 cosmCladocera Chydorus sphaericus (OF Muumlller 1785) 4 cosm species groupCladocera Daphnia laevis Birge 1879 278 tropicopol North South America

and Africa onlyCladocera Dunhevedia serrata Daday 1898 45 tropicopolCladocera Euryalona orientalis (Daday 1898) 4 tropicopolCladocera Ilyocryptus spinifer Herrik 1882 58 tropicopolCladocera Karualona socotrana Dumont amp Silva-Briano 2000 5 afrotrop SokotraCladocera Kurzia longirostris (Daday 1898) 5 circumtrop eastern hemisphereCladocera Leberis diaphanus (King 1853) 2 circumtrop eastern hemisphereCladocera Macrothrix spinosa King 1853 345 tropicopolCladocera Moinodaphnia macleayi (King 1853) 4 torpicopolCladocera Notoalona globulosa (Daday 1898) 5 circumtropOstracoda Cryptocandona sp 2 palaearct introducedOstracoda Cypretta cf seurati Gauthier 1929 4 palaerct neotrop orient pacificOstracoda Cypria cf lenticularis GW Muumlller 1898 7 afrotropOstracoda Cyprididae Gen sp 8Ostracoda Gomphocythere obtusata Sars 1910 78 afrotropOstracoda Nealecypris obtusa (Klie 1933) 14 afrotropOstracoda Potamocypris sp 2Ostracoda Zonocypris costata (Vaacutevra 1897) 27 palaearct afrotropAcari Tactocepheus cf velatus (Michael 1880) 2 cosmAcari Trhypochtoniellus longisetus (Berlese) f setosus Willmann 1928 2 cosmAcari Punctoribates sp 3

(Continued)



INTRODUCTION

The island of Madagascar is among the tophotspots for biodiversity conservation worldwide(Myers et al 2000 Ganzhorn et al 2008 Vieiteset al 2009) More than 90 of all plants 100 ofnative amphibian and mammal species 92 ofreptiles and 44 of birds occur nowhere else Thishigh level of endemism predominantly results fromradiation of African founder individuals arrivingduring the Cenozoic (655Ma to present) and fromrelict species radiating after the island separatedfrom the African mainland (183ndash158Ma)Antarctica (130Ma) and India (96ndash65Ma Venceset al 2009)

Besides an extraordinary diversity in terrestrialorganisms Madagascar has also been recognizedas a global hotspot of freshwater biodiversity(Groombridge and Jenkins 1998) In the last twodecades of the 20th century the number of newlydescribed species of aquatic insects and fish hadincreased exponentially (Benstead et al 2003)However owing to incomplete species inventoriesand taxonomic difficulties the degree of endemismin small planktonic and benthic organisms isdifficult to assess There is evidence for endemismespecially for copepods (Dussart 1982) but alsofor some freshwater algae (eg desmids (West andWest 1895 Bourrelly and Couteacute 1991 Coesel2002) and benthic diatoms (Metzeltin and Lange ndash

Bertalot 2002))Taxonomic research on Malagasy freshwater

algae and micrometazoa is still scarce There areno systematic species inventories available andcurrent knowledge is based on random samplesmostly covering specific groups of organisms fromdifferent habitats First surveys of the Malagasyfreshwater algal flora were provided by West andWest (1895) Fritsch (1914) Manguin (1941)Bourrelly and Leboime (1946) and Bourrelly andManguin (1949) More detailed algologicalinvestigations are available for diatoms (Spauldingand Kociolek 1998a b Metzeltin and Lange ndash

Bertalot 2002) desmids (Bourrelly and Couteacute1991 Coesel 2002) and chrysophytes (Hansen1996) Taxonomic research on micrometazoa wasconducted predominantly until the 1960s(DeGuerne and Richard 1893 Brehm 19301948 Kiefer 1930) Most studies were based onsamples taken by the director of the Institut desrecherches scientifiques de Madagascar RenauldPaulian who sent the material to varioustaxonomists (Rotifera Berzins 1960 1973 1982

Harpacticoida Chappuis 1952 1954 1956Calanoida Brehm 1951 1952a b c 1953 1954Cyclopoida Lindberg 1951a b 1952 1953 Kiefer1952 1954 1955 Cladocera Brehm 1953) Brehm(1960) gave an account of freshwater Crustaceacollected during the Austrian Madagascarexpedition in 1958 After this period of intensivetaxonomic work on freshwater micrometazoa fewpublications appeared (Dussart 1982 Segers 1992Fiers 2002)

Madagascarrsquos biological richness is threatened bymassive habitat destruction Forty per cent ofrainforest cover was lost between 1950 and 2000(Harper et al 2007 Wikipedia 2012)Deforestation currently continues at a rate ofabout 05 per year (57 000 ha Mongabay 2011)Although the country possesses 46 legallyprotected areas covering almost 17 millionhectares (289 of total territory) habitatdestruction still proceeds at an alarming pace Apeak in illegal logging was observed during thepolitical turmoil in 2009 Schuurman and Lowry(2009) spoke of a lsquoMadagascar rosewoodmassacrersquo because 625 containers of hardwoodworth approximately US $ 130 million illegally leftthe country in 2009 Most of the trees were felledin protected mountain reserves in NorthernMadagascar (Marojeji Masoala and AmberMountains) During sampling in May 2009 theillegal harvest of rosewood and palisander(Dalbergia spp) was witnessed on several occasions

Deforestation not only changes terrestrialhabitats but causes increased sediment deliveryand nutrient input into rivers and lakes and hencealters physico-chemical conditions speciescomposition and trophic interactions in aquatichabitats as well (Benstead et al 2003) Loss offorest cover has already caused a shift frompersistent to intermittent flow in the streamsdraining the northern and eastern slopes of theAmber Mountains and the Tsaratanana Massif(Raxworthy et al 2008)

Following habitat destruction invasive speciesare regarded as the second leading cause of speciesextinction and endangerment worldwide (Salaet al 2000) A number of alien fish species havebeen stocked in pristine Malagasy fresh waterscausing severe changes to entire ecosystems andbeing detrimental to various groups of animals(Canonico et al 2005) For instance there is astrong correlation between the introduction ofexotic fish and the decline of native fish inMadagascar (Reinthal and Stiassny 1991 Sparks









STUDY AREA






METHODS




RESULTS




























Species assemblages


able

1Description

oflakesin

theAmberMou

ntains

Nationa

lPark

Mad

agascar(indicatesna

tive

fish

species)

Lak

eNum

ber

Malagasy

Nam

eForeign

Nam

eSam

pling

Date

Altitude

(m)

Max

Length

(m)

Max

Depth

(m)

Tem

p(C)

Cond

(mScm

-1)

pHOxy

gen

(mgL-1)

Oxy

gen

(sat)

Fish

1Mah

asarika

PetitLac

16052009

1060

280

48

246

28928

824

115

Tila

piarendalli

Cyprinuscarpio

Micropterus

salm

oides

Gam

busiaaffinis

2Malio

Grand

Lac

16052009

1335

360

40

206

31841

797

105

CyprinuscarpioM

icropterus

salm

oides

3Taran

taMau

dit

18052009

1250

720

09

198

56822

809

103

CyprinuscarpioM

icropterus

salm

oides

4Mah

ery

Mah

ery

19052009

364

765

Shallow

261

218

687

--

Tila

piarendalli

Paratila

piasp

5Antagna

voely

Sacreacutesm

all

20052009

348

190

25

259

202

716

--

Tila

piarendalli

Paratila

piasp

Ptychochrom

issp

6Antagna

voSa

creacute

20052009

367

1660

261

231

874

Tila

piarendalli

Paratila

piasp

Ptychochrom

issp

7Man

onja

Texier

22052009

1034

630

25227

61901

872

113

nointrod

uced

fishe

els

8Fan

tany

Fan

tany

23052009

810

780

32254

75906

801

108

nointrod

uced

fishe

els













DISCUSSION




















ACKNOWLEDGEMENTS


REFERENCES






















































































APPENDIX



Number Biogeography




(Continues)



Table 1 Continued









(Continues)

(Continued)



Table 1 Continued



West 19044 cosm





(Continues)

(Continued)



Table 1 Continued





(Continues)

(Continued)



Table 1 Continued






(Continued)









STUDY AREA






METHODS




RESULTS




























Species assemblages


able

1Description

oflakesin

theAmberMou

ntains

Nationa

lPark

Mad

agascar(indicatesna

tive

fish

species)

Lak

eNum

ber

Malagasy

Nam

eForeign

Nam

eSam

pling

Date

Altitude

(m)

Max

Length

(m)

Max

Depth

(m)

Tem

p(C)

Cond

(mScm

-1)

pHOxy

gen

(mgL-1)

Oxy

gen

(sat)

Fish

1Mah

asarika

PetitLac

16052009

1060

280

48

246

28928

824

115

Tila

piarendalli

Cyprinuscarpio

Micropterus

salm

oides

Gam

busiaaffinis

2Malio

Grand

Lac

16052009

1335

360

40

206

31841

797

105

CyprinuscarpioM

icropterus

salm

oides

3Taran

taMau

dit

18052009

1250

720

09

198

56822

809

103

CyprinuscarpioM

icropterus

salm

oides

4Mah

ery

Mah

ery

19052009

364

765

Shallow

261

218

687

--

Tila

piarendalli

Paratila

piasp

5Antagna

voely

Sacreacutesm

all

20052009

348

190

25

259

202

716

--

Tila

piarendalli

Paratila

piasp

Ptychochrom

issp

6Antagna

voSa

creacute

20052009

367

1660

261

231

874

Tila

piarendalli

Paratila

piasp

Ptychochrom

issp

7Man

onja

Texier

22052009

1034

630

25227

61901

872

113

nointrod

uced

fishe

els

8Fan

tany

Fan

tany

23052009

810

780

32254

75906

801

108

nointrod

uced

fishe

els













DISCUSSION




















ACKNOWLEDGEMENTS


REFERENCES






















































































APPENDIX



Number Biogeography




(Continues)



Table 1 Continued









(Continues)

(Continued)



Table 1 Continued



West 19044 cosm





(Continues)

(Continued)



Table 1 Continued





(Continues)

(Continued)



Table 1 Continued






(Continued)



METHODS




RESULTS




























Species assemblages


able

1Description

oflakesin

theAmberMou

ntains

Nationa

lPark

Mad

agascar(indicatesna

tive

fish

species)

Lak

eNum

ber

Malagasy

Nam

eForeign

Nam

eSam

pling

Date

Altitude

(m)

Max

Length

(m)

Max

Depth

(m)

Tem

p(C)

Cond

(mScm

-1)

pHOxy

gen

(mgL-1)

Oxy

gen

(sat)

Fish

1Mah

asarika

PetitLac

16052009

1060

280

48

246

28928

824

115

Tila

piarendalli

Cyprinuscarpio

Micropterus

salm

oides

Gam

busiaaffinis

2Malio

Grand

Lac

16052009

1335

360

40

206

31841

797

105

CyprinuscarpioM

icropterus

salm

oides

3Taran

taMau

dit

18052009

1250

720

09

198

56822

809

103

CyprinuscarpioM

icropterus

salm

oides

4Mah

ery

Mah

ery

19052009

364

765

Shallow

261

218

687

--

Tila

piarendalli

Paratila

piasp

5Antagna

voely

Sacreacutesm

all

20052009

348

190

25

259

202

716

--

Tila

piarendalli

Paratila

piasp

Ptychochrom

issp

6Antagna

voSa

creacute

20052009

367

1660

261

231

874

Tila

piarendalli

Paratila

piasp

Ptychochrom

issp

7Man

onja

Texier

22052009

1034

630

25227

61901

872

113

nointrod

uced

fishe

els

8Fan

tany

Fan

tany

23052009

810

780

32254

75906

801

108

nointrod

uced

fishe

els













DISCUSSION




















ACKNOWLEDGEMENTS


REFERENCES






















































































APPENDIX



Number Biogeography




(Continues)



Table 1 Continued









(Continues)

(Continued)



Table 1 Continued



West 19044 cosm





(Continues)

(Continued)



Table 1 Continued





(Continues)

(Continued)



Table 1 Continued






(Continued)






















Species assemblages


able

1Description

oflakesin

theAmberMou

ntains

Nationa

lPark

Mad

agascar(indicatesna

tive

fish

species)

Lak

eNum

ber

Malagasy

Nam

eForeign

Nam

eSam

pling

Date

Altitude

(m)

Max

Length

(m)

Max

Depth

(m)

Tem

p(C)

Cond

(mScm

-1)

pHOxy

gen

(mgL-1)

Oxy

gen

(sat)

Fish

1Mah

asarika

PetitLac

16052009

1060

280

48

246

28928

824

115

Tila

piarendalli

Cyprinuscarpio

Micropterus

salm

oides

Gam

busiaaffinis

2Malio

Grand

Lac

16052009

1335

360

40

206

31841

797

105

CyprinuscarpioM

icropterus

salm

oides

3Taran

taMau

dit

18052009

1250

720

09

198

56822

809

103

CyprinuscarpioM

icropterus

salm

oides

4Mah

ery

Mah

ery

19052009

364

765

Shallow

261

218

687

--

Tila

piarendalli

Paratila

piasp

5Antagna

voely

Sacreacutesm

all

20052009

348

190

25

259

202

716

--

Tila

piarendalli

Paratila

piasp

Ptychochrom

issp

6Antagna

voSa

creacute

20052009

367

1660

261

231

874

Tila

piarendalli

Paratila

piasp

Ptychochrom

issp

7Man

onja

Texier

22052009

1034

630

25227

61901

872

113

nointrod

uced

fishe

els

8Fan

tany

Fan

tany

23052009

810

780

32254

75906

801

108

nointrod

uced

fishe

els













DISCUSSION




















ACKNOWLEDGEMENTS


REFERENCES






















































































APPENDIX



Number Biogeography




(Continues)



Table 1 Continued









(Continues)

(Continued)



Table 1 Continued



West 19044 cosm





(Continues)

(Continued)



Table 1 Continued





(Continues)

(Continued)



Table 1 Continued






(Continued)













DISCUSSION




















ACKNOWLEDGEMENTS


REFERENCES






















































































APPENDIX



Number Biogeography




(Continues)



Table 1 Continued









(Continues)

(Continued)



Table 1 Continued



West 19044 cosm





(Continues)

(Continued)



Table 1 Continued





(Continues)

(Continued)



Table 1 Continued






(Continued)















ACKNOWLEDGEMENTS


REFERENCES






















































































APPENDIX



Number Biogeography




(Continues)



Table 1 Continued









(Continues)

(Continued)



Table 1 Continued



West 19044 cosm





(Continues)

(Continued)



Table 1 Continued





(Continues)

(Continued)



Table 1 Continued






(Continued)





















































APPENDIX



Number Biogeography




(Continues)



Table 1 Continued









(Continues)

(Continued)



Table 1 Continued



West 19044 cosm





(Continues)

(Continued)



Table 1 Continued





(Continues)

(Continued)



Table 1 Continued






(Continued)



Table 1 Continued









(Continues)

(Continued)



Table 1 Continued



West 19044 cosm





(Continues)

(Continued)



Table 1 Continued





(Continues)

(Continued)



Table 1 Continued






(Continued)



Table 1 Continued



West 19044 cosm





(Continues)

(Continued)



Table 1 Continued





(Continues)

(Continued)



Table 1 Continued






(Continued)



Table 1 Continued





(Continues)

(Continued)



Table 1 Continued






(Continued)



Table 1 Continued






(Continued)



on the brink - investigating biodiversity in endangered crater lakes of the amber mountains national...

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