ECOGRAPHY 21: 371-382. Copenhagfn 1998

Biogeographical regions of the Iberian peninsula based onfreshwater fish and amphibian distributions

J. Mario Vargas, Raimundo Real and Jose C. Guerrero

Vargas, J. M., Real, R. and Guerrero. J. C. 1998. Biogeogt-aphical regions oftheIberian peninsula based on freshwater fish and amphibian distributions. - Bcographv21; 371 382.

We classified the main Iberian river basins based on the presence and absence offreshwater fishes and amphibians. Tor both taxonomie groups we analysed three datasets; 1) endemic species only, to search for biotic boundaries related to historicalevents, 2) indigenous species, which include endemic ones, to search for bioticboundaries related to ecological factors, 3) indigenous and well-established intro-duced species, to assess the inlluence of man in the current biogeographieal patternso\' lishes and amphibians. We used both phenetic and cladistic methods, followed bya consensus analysis to provide an overall biogeographical pattern. Based on all fishdistributions, the Iberian Peninsula is divided into three biogeographical regions:Cantabrian, Atlantic and Mediterranean, No boundary existed between theCantabrian and Atlantic regions when only indigenous fish species were considered.This suggests that this boundary has been induced by man, probably through thedifferentia! introduction of hsh species into reservoirs at one or other side of theboundary. Run-off and the size of the river basins are the environmental factors thatdistinguished the Atlantic and Mediterranean regions. However, regionalizationbased only on endemic freshwater fishes showed a la^titudinal pattern thai agrees withthe paleogeographic events of the Upper Oligocene-Lower Miocene period. Bycontrast, one northem and one southern region were distinguished based on allamphibian distributions and on indigenous amphibians only, which suggests thathuman activity has not significantly affected the overall biogeographical pattern ofamphibians in the Iberian Peninsula. Interannual predictability of precipitation bestaccounts for this regionalization. Based on endemic amphibians, the Iberian Penin-sula is divided into three regions that closely resemble the three separate land areasof the Upper Eocene-Lower Oligocene period. The consensus between the biogeo-graphical regions based on lishes and amphibians yields five pairs of basins. Geolog-ical origin of the basins seems to better explain the consensus between thebiogeographical patterns of fishes and amphibians, whereas ecological factors proba-bly contribute to the differences between them.

J. M. Vargas tjmvy(a),wna,c.s). R. Real and J. C. Guerrero. DepI of Animal Biolot^v.Fac. ('/ ScieiJcex, Univ. of Malaga. E-29071 Malaga. Spain.

Ft-eshwater lishes and amphibians are charactefizeLl The tt.se of two or more distantly related taxa forhistoncally by their relative antiqttity. and ecologically dctnarcating biogeogfaphical tegions increases the reli-by tlieir poor dispersal ability and the ecophysiologi- ability o\' the regionalization. because the greater thecal dependence on freshwater. Such similarities might number of animal and plant groups that reveal lliemake freshwater lishes and amphibians suitable for satne patterns, the more robust these patterns are.itivestigatiiig the possible existence of shared biogeo- However, few authors have analysed the distributiongraphical regions. patterns shared by different groups o\~ orgaiiisrns

Accepted 20 October 1997

Copyright ••(" HCOGRAi'HY 1998ISSN 0906-7590Printed in Ireland all rights reserved

ECOCIUAI'MY 21:4 (I y9K)

(Holloway and Jardiiic 1968, Endler 1982. Griiluini

1990).In recent years quantitalive methods have increased

the ri^or of elucidating biogeographical regions (Crov-dlo 1981, Brown and Gibson 19H3. Birks 1987). andquantitative nuiltivariate techniques (Gauch 1982) havebeen used to Iind similar groups of Operational Geo-graphic Units (OGUs sensu Crovello 198!). These tech-niques examine the data in two different andcomplementary ways: ordination and classification: de-pending on whetlicr the areas are placed a priori intheoretically continuous or discontinuous sequences(McCoy et al. 1986). The analytical technique of McCoyet al. (1986) and modified by Real et al. (1992b) andMarqucz et al. (1997) enables the researcher to testwhether species are distributed independently of oneanother in a random continuous sequence or. alterna-tively, species are nonrandomly distributed, with theircombined ranges demarcating biogeographical regions.In addition, discriminant functions may be used tointerpret the biotic regions in terms of environmentalvariables (see ter Braak I9S6, Myklestad and Birks

1993).The Iberian Peninsula hosts a considerable diversity

and heterogeneity in its freshwater tish (Doadrio 1988)and herpetological faunas (Schall and Pianka 1977.Busack and .laksic 1982). This region is typified by itsphysiographical heterogeneity, its geographical situationin southwestern Europe near Africa, and the fact that itserved as a refuge during the Quaternary Glacial Age(de Lattin 1967). At present, the Strait of Gibraltar andthe Pyrenees act as barriers separating the IberianPeninsula from the larger European and African re-gions. As a result, the Iberian Peninsula constitutes adistinctive biogeographical entity of 580 000 km- withinthe western Palaearctic, for freshwater fishes (Doadrio1988) and amphibians (Real et al. 1992a).

The peninsula comprises 11 main river basins (Fig. 1).River basins are infomiative OGUs because they tbrmnatural geographical units with well-defmcd borders(Birks 1987). and they are often used in biogeographicalstudies of freshwater organisms (see, e.g. Sepkoski andRex 1974. Matthews and Robison 1988, Hugueny andLeveque 1994). River basins have ecological value be-cause they are self contained with respect to the nettransport of nutrients (Margaief 1980). their externalboundaries may reduce the movements from one basinto another, and the mountains thai form the basinsaffect both rains and run-offs, the main factors respon-sible for doods (Real et al. 1993).

In their present form, the Spanish river basins datetrom the Quaternary, and before this time large endor-rheic basins occupied the entire Iberian Peninsula(Doadrio 1988). The orogeny of the Pyrenees began inthe Lower Eocene, in an east-west orientation, and theytook on their current tbrm in the Upper Oligocene-Lowcr Miocene. During the latter period, the Iberian

Mountain System emerged (Lopez-Martinez 1989). TheFbro-Eastern Pyrenees pair of basins (Fig. 1) became thelluvial network between the Pyrenees and the IberianMountain System. The Cantabrian Range rose in awest-east direction in the Upper Oligocene-MiddleMiocene, and the North-Mifio pair of basins tbrmed inits north-western slope. The Duero and Tajo basins wereformed as single basins in the Lower-Middle Miocenealthough they began to separate during the formation olthe Central System in the Lower Oligocene. The endor-rheic basin in the southern subplaieau drained into theAtlantic through the Guadiana and the Guadalquivir(Upper Miocene), which only were partially separated intheir middle and upper sections by the base of the SierraMorena. The South-Segura pair of basins (Fig. I) devel-oped into the south-eastern drainage system in the Beticrange (RogI and Steininger 1983), after tlie breakingaway of the Bctic-Riff Massif when the Strait of Gibral-tar was formed (Upper Miocene-Pliocene). The Jucarbasin formed the opening of the southern subplateau tothe Mediterranean in the Pliocene.

Most of the presenl-day freshwater fish and amphib-ian species were present during the Upper Oligocenc-Miocene (Doadrio 1990). when the Iberian Peninsulawas separate from Africa and the Pyrenees were alreadyan effective barrier (Lopez-Martine? 1989). As a conse-quence, both types of fauna display a high degree ofendemicity.

Herein we analyse the biogeographica! patterns of thefreshwater fishes and amphibians of the Iberian Penin-sula, searching for biotic boundaries between theIberian river basins and their historical, ecological orman induced causes. This comparative approach en-ables an evaluation of whether the two groups: a) sharebiogeographical regions, b) have different patterns ofbiotic regionalization. or c) exhibit partially shared andpartially separate patterns.

Material and methods

The geographical units and the speciesdistributions

The distributions of freshwater fish species (n - 51) mthe Iberian Peninsula were taken from Doadrio et al.(1991), and the distributions of amphibian species (n =25) from Martinez-Rica (1989) and Pleguezuelos (1997).We included as freshwater fishes the diadiomous spe-cies, but excluded from the analysis the estuarine orperipheral species that do not maintain permanent pop-ulations in freshwater. No new data modify the distribu-tion of fishes in the river basins reported by Doadrio etal. (1991), However, two modifications were made toamphibian data: a) Tritiiriis nminwratus (Latreille) wastreated as two species. T. mainwraius and T. py^niacus.although it is not clear whether they represent differentspecies""or subspecies (Garcia-Paris et al. 1993), b) Rana

372LX'Of.RM'IIY

F i g . I. I b c r i i m I'cLiirisiiia d i v i d e d i n l o i ls I 1 m a i n r i v e r b a s i n s .

pyycnaica Scrra-Cobo has not been considered becauseIt is ii too recently described species whose status anddistribution is insufficiently known {Serra-Cobo 1993).

We analysed three datasets for each taxonomicgroup: I) endemic species only, lo search for historiealpatterns: 2) indigenous species, which included endemicspecies, to search for ecological patterns; howevei". thedistributions ol' some indigenous species within theIberian Peninsula may have been altered by speciesintroduced by humans, many of which are known loplay a well-defined ecological role (Hernando andSoriguer 1992), so that we used dataset 3) indigenousand well-established introduced species, to assess theinfluence of man in the current distribution patterns oftishes and amphibians.

Classification methods

We used cluster analysis and cladistic analysis to clas-sify rivei- basins according to presence or absence ofspecies, as they represent two diftc-rcnt but complemen-tary approaches. The cluster analysis enabled us togroup the OGUs, which are delimited only by easilyrecognizable geographical boundaries, into Operational

Biogeographic Units (OBUs). which are delimited bybiotic boundaries. The cladistic analysis enabled us toobtain the most parsimonious relationship betweenbasins according to their shared presences of species(Rosen 1984. Myers J991).

In the cluster analysis, we applied the UPGMA(I'nweighled Pair-Group Melhod using arithmetic Av-erages) algorithm (.Sneath and Sokal 1973) to two dif-ferent similarity matrices based on all pairwisecomparisons of river basins, a Jaccard (191)! ) matrixand a Baroni-Urbani and Buser (1976) matrix, becausethe probabilities associated with the values of thesecoefficients can be calculated (Baroni-Urbani and Buser1976, Real and Vargas 1996). .laccard's coefficient doesnot take into account double absences: as a result, thesimilarity values between pairs of OGUs do not dependon the presence of other species in other OGUs. Inaddition, each Jaccard's value is related to a differentnumber of species (N), those shared by the two basinscompared, and therefore a more extreme value may benon significant because of a low N, while a moremoderate value with a higher N could be statisticallysignificant. In contrast. Baroni-Urbani and Buser's co-efficient considers double absences, and so, the similar-ity values between the pairs of OGUs depend on otherspecies that may only be present in other OGUs. Withthis index a more extreme value is always more signifi-cant than a more moderate ont, because the number ofspecies (N) related to these values is the same, that is.all the species present in the study area. The similarityvalues were transformed into three classes identified by'' + ". •'-"" and "()•" signs, according to whether thevalues were significantly higher. lower or the same asexpected at random; to do so we used the table inBaroni-Urbani and Buser (1976) for the Baroni-Urbaniand Buser coefficient, and the formulae in Real andVargas (1996) for the Jaccard coefficient.

We modified the method of McCoy et al. (1986) totest lor the existence of significantly strong or weakbiotic boundaries as follows. For each dentiiogramnode we established a submatrix of significant similari-ties that only included the basins involved in the node.This submatrix was divided into three zones: zone Aand zone B, which corresponded to each group ofbasins separated by the node: and zone A x B, corre-sponding to the intersection between both zones. Theparameters DW (A x A) and DW (B x B) measure theextent to which the similarities that are higher thanexpected ( +) tend to be in zones A and B. but not inA X B (see McCoy et al. 1986). The parameter DS givesa measure of whether the similarities lower than ex-peeted ( - ) tend to be located in A x B. but not in A orB. The statistical significance of the node was assessedby a G lest of independence of the distribution of thesigns " + ". ••-"" and -'O" in the three zones of thesubmatrix. giving the parameters GW, for weakboundaries, and GS, for strong boundaries. If similari-

373

ties higlief than expected ( + ) tire non-rLindomlygrouped in zones A and B, but not in A x B, thenthere is at least a weiik boundary between bothgroups of OGLis. If similarities lower than expected( - ) are non-randomly grouped in A x B. but not inA or B. then a strong boundary exists between ihc

groups.In the cladislic analysis only the syuapomorphic

species are informative, that is. those species presentin tnore than om river basin but not present in allbasins. We used tlie Phylogenetic Analysis Using Par-simony (PAUP) software (Swoftord 1993), selectingFitches parsimony method (Fitch 1971), because ilconsiders the characters as reversible, and the Branch-and-Bound algorithm, that gives a mathematical guar-antee of finding all optimal solutions. The bootstrapoption based on 100 replicates was applied whenmore than one most parsimonious solution was ob-tained. We rooted the trees using the oulgroupmethod, the outgroup being all absences, and carriedout the character state optimization according to theAccelerated Transformation (ACCTRAN) option.The consensus between the results obtained with UP-GMA and PAUP was carried out by Brooks's parsi-mony analysis (BPA) (Brooks 1981. 19S6), and abootstrap analysis was performed on the resultingcomponents matrix.

Table I. Environmental variables used in tlie analysis

Geographical variables Code

Average geographie longitude'Latitude'Elevation'Hlevatiou range'Distance from the coast'Distance Ironi tlie isthmus'Area'

(LO)(LA)(E)(BR)(DC)(DI)(A)

Climatic variables Code

Mean aunual precipitation"'Absolute maximum precipitation

recorded in 24 h"Relative maximum precipitalioriMean annual number of days with

precipitation'Mean relative humidity at 07.00

for January^Mean auuual temperature-Differences between the mean

temperatures of January and July-Annual mean hours of sunshine-Global solar radiation in

kWh m • =/day^Potential evapotranspiration-Interamiual pluviometric irregularity'Run-ofCActual evapotranspiration''

(P)

(MP)(RMP)

(DIM

(H)(T)

(TR)(HSI

(SR)(PET)(PU(RO)(AHT)

Sources' l)maps 1:800 000 from Spanish Military GeographieService, 2) Font (1983). 3) Montero dc Burgos and Goiizalc/-Rebollar (1974). 4) Auon, (1979).

Environmental variables

We obtained data of 20 variables that estimate the maingeographical and climatic features of the basins (TableI), with the aim of searching for the environmentalfactors that best distinguished the regions obtained.The relative maximum precipitation (RMP) was derivedfrom the mean annual precipitation (P) and the maxi-mmn precipitation recorded in 24 h (MP) (RMP = MP/P) and represents an index of the potential severity ofHoods (Real ct al. 1993), The interannual pluviometricirregularity (PI) is defined as the coefficient of variationin annual precipitation, and gives a measure ol theinterannual predictability of precipitation.

We characterized the OBUs identified by clusteranalysis in environmental terms by means ol" stepwisenon-standardized canonical discriminant functions, us-ing SPSS-X (1988) software. We halted the discrimi-nant analysis when 1OO' > of the OGUs had beencorrectly classified into the corresponding OBU. usingthe above variables as discriminants. When there weremore than two OBUs in a regionalization pattern, ahierarchical relationship was established between themby PAUP. and then we applied the discriminant anal-ysis to the branches separated at each node of theresulting tree.

Results

Iberian biogeographical regions based onfreshwater lisbes

AH spceie.sUsing .laccard's coefficient and UPGMA. we only de-tected one weak boundary based on all freshwaterfishes, separating two OBUs: the group of Atlanticbasins 3. 6, 8 and 9: and the group of Mediterraneanbasins 4. 5 and 7. The two Cantabrian basins (I and2) and the south-eastern basins (10 and II) neithermake up independent regions, nor do they significantlyfonn part of either of the aforementioned OBUs (Fig.2a. Table 2). They support fewer species than the twoOBUs identified (see Appendix 1) and thus their bio-geographical identity is not significantly defined byJaccard's coefficient, which considers similarity basedon shared species present in the basins, but not shared

abscnces-Using Baroni-Urbani and Buser^ index, we detected

two weak boundaries that separate three OBUs in theIberian Peninsula; each OBU was related to a differentwatershed: Cantabrian, Atlantic and Mediterranean(Fig. 2b, Table 2). With this index all the OGUs wereclassified into one of the OBUs. This indexsignificantly discriminates among basuis 1, 2. 10, and11. because it considers double presences and doubleabsences.

374ECOCiRAPHV 2t;4

l.'sing Barcmi-Urbani and Biisci's index, we detectedtwo weak boundaries that separate three OBUs in theIberian Peninstila; each OBU was fckuod to a differentwatershed; Caiitabriaii, Atlantic and Mediterranean(Fig. 2b. Table 2). With this index all the OGUs wereclassified into one of tlie OBLis. This index signifi-cantly discriminalcs among basins 1. 2. 10. and 1!,becatise it considers double presences and double ab-sences.

Using PAUP (Fig. 2c), the Cantabrian basins (I and2) form one group, the Atlantic basins (3. 6. 8 and 9)another, and the three northern Mediterranean basins(4. 5 iind 7) a third, whereas the south-eastern ri\erbasins (10 and II) segregate as individual basins. Itmay be thai the low number of synapomorphic speciespresent in basin 11 (only 8 species) and the fact thatthey are present in many basins prevent basin 11 frombeing associated with other basins. The same mayoceur to a lesser extent for basin 10.

The consensus between PAUP and the two treesobtained using UPGMA is shown in Kig. 2d. Thisconsensus tree is similar to that produced by Jaeeard'sindex, but it clearly favors the existenee of aCantabrian region, since the frequency of occurrencein the bootstrap analysis is 96'K> for the Cantabrianbasin group and l()D'l/ii for the remaining basins. Theconsensus also iavors the existenee of a region com-prising tlie Atlantic basins 3. b. 8 and 9, and one oihcrregion tliat includes the Mediterranean basins 4. 5 and7. Only the group formed by basins 10 and II has arelatively low level of eonsensus. with a 74'%i frequencyof oceiurence. and only 77"'<i for ihe other branch ofthe fork. Thus the position of these two basins is moreambiguous.

These results suggest the existence of three regionsfor freshwater fishes. Cantabrian. Atlantic, and Med-iterranean, when double absences are considered, asdetected with Baroni-Urbani and Buser's index. This isthe only index that unambiguously classifies all basins,whereas the position of tiie southern Mediterraneanbasins (10 and 11) remains unclear with Jaccard's in-dex and PAUP.

Indigenous speciesWhen the UPGMA method was applied to indigenousfreshwater fishes (n = 37). no significant boundary wasdetected with .laccard's intlex (Fig. 2e. Table 2).whereas Baroni-l.lrbani and Buser's index showed theexistence of a weak biotic boundary separating theMediterranean region from the Cantabro-AtUintic re-gion (I*ig. 2f, Table 2). This boundary results from theabsence of species from either side of the bouiuiary.because tinly the index that takes double absences intoaccount was able to detect it. The tree obtained withPAUP (Fig. 2g) confirms Ihe Cantabro-Atlantic regionfomied by basins 1. 2. 3. 6, 8 and 9. but not iheMediterranean region, because the pair of basins [()

2 isJS 2

o o

^ <

~, 3SO

r i -3

f , r~; o— ni i-i

"•. r-i cs r-i OSn r^ f l f, —ID i~i i-o

I

— ^ — ^ 00 —Ir~- ^ — O r<-. lyjO —^ 00 r>i — 1^

o c; c: o c;

^ iO r-- r i r-- (--I — csI I . I I

oc 00 x: ocoooc c o o

l.COC.RAI'UY 1\ 375

all especies

0,1 0,2 0,3 Q.4 05 0.6 07 08 0,9

indigenous speciesDO 0,1 02 0.3 0 / 0.5 Ofi 0.7 0,B 0,9

endemic especiesoo 01 02 03 04 05 06 07 08

0.0 0.1 0£ 0.3 04 05 OS 07 08 09 10 0.0 0.1 0.2 0.3 0.1 0.5 06 0.7 OS 09 10 0.0 0 1 0.2 0.3 0.4 0.5 06 0.7 O.S 0.9 1.0

Fig. 2. Classificalion ol' ilicIbcriiin river basinsaccording lo fresliwiiler fislidislribulion for all species(tirs! column), indigenousspecies (second Lokimn) andendemic species (lliirdcolunm). using J;iecard'sindex (first row),llaroni-Urbani and Buser'sindex (second row), PAUP(third row) itiid bootstrapon BPA (lomlh row). W:Weak biodc boundary; S:Strong biotic boundary.** = p<().()l.

cind 11 segregate Ironi the other basitis in theregion.

The consensus between the three classification tncth-ods based on indigenous fishes (Fig. 2h) confirms theCantabro-Atlantic region formed by basins I, 2, 3. 6, 8and 9. wilh a frequency of occurrence in ihe bootstrapanalysis of 9rVii, and a Mediterranean regioti compris-ing Ihe basins 4, 5 atid 7, with 95'Vli of occurrence. Tbeposition of basins H) and 11 is not clearly supported inthe consensus tree, because the frequency of oceurrenceof the group formed by the rest of the basins is only73'! !. So. the classification with liaroni-Urbatii andBuser's index is again ihe only one that unatnbiguouslyclassifies all Iberian basins in one of the two OBUsobtained.

Endemic speciesWhen only endemic fish were considered, both Jac-card's and Baroni-Urbani and Buser's indices separatedthe most southerly river basins (8, 9. 10 and 11) frotnthe others (Fig. 2i, 2j) but only Baroni-Urbani andBuser's index detected a strona biotic boundary be-

tween the two groups of basins (Table 2). This patternis also the tnost parsimonious obtained using PAUP.After applying liPA, the northern region has 99"A- ofoccurrence and the southern region 94'/ . (Fig. 21), andso both OBUs are strongly supported.

Division of the Iberian Peninsula for amphibians

All amphibiansThe two binary indices produce the same regionaliza-tion of the Iberian Peninsula for all amphibians, basedon a weak boundary that separates two OBUs latitu-dinally (Fig. 3a, 3b. Table 3). Therefore, this pattern issupported when double absences are considered, andalso when only the species shared between the basinsare taken into account. In the single most parsirno-nious tree retained by PAUP the basins are chained(Fig. 3c), which is charactetnstic of weak separationbetween the basins. It is worth noting that 8 out of 25species are noninformative when using PAUP, becausethey are present either in all basins or in only one

376 E t X K H i A l ' U Y 2\A (!'»•>!<)

Fig. ?•. Classiticaiion of llicIberian river basinsaccoidirig (o amphibian(.iislribijlion ibr ali species(lirsl coiumnl. indigenousspecies (second coltimn) andendemic species (ihirdcolumn), using Jaticard'sindex (llrsl row),Baroni-Urb;ui! and Buser'sindex (second row), PAUP(third row) luid boolstrapon BI'A (Iburtli row). W:Weak biolic boundary; S:Slroni; biolic boundary.* = p<l),U5: ** = b

all especies indigenous species endemic especies0 6 0 7 0 S 0 9 1 . 0 0 0 0 ] 0 2 0 1 0 a 0 5 0 6 0 7 0 B 0 9 1.0 0 0 0 1 0 2 0 3 0 - 1 0 5 0 6 0 7 0 8 0 9 1 0

O . Q O 1 O 2 O J O 4 O 5 O 6 0 7 D 8 O 9 10 0 0 0 1 0 J 0 3 D 4 0 5 0 6 0 7 0 8 C 9 1 0 0 0 0 1 0 2 0 3 0 4 0 5 0 6 0 7 0 8 0 9 1 0

97

1-}

56 rL

99

rL

j ^

78 [

49

7

56 r

99

rL

78 1

JIL|

—1

78 r

1

1

75

699S j

50 f~^^±-

180 m -

^ 3

to

J 11 ^

( \

7J '01 11

basin. This problem atTects basins Id, Ii and 7. andreduces the species list in half. In iidtiition. theinformative species of basin 7 are widespread (seeAppendix 1) and so this basin is not grouped with anyspecific basin.

Consensus between PAUP and the two treesobtained from UPGMA is shown in Fig. 3d. Theregionalization is similar to that obtained withJaecard's and Baroni-Urbani and Buser's indices,although basin 7 is not clearly related to either of thetwo regions. So the basins of Gnadiana (8),Guadalquivir (9). Segura (10) and the smaller southernSpanish river basins (II) seem to form a southern OBUfor amphibians, whereas the rest of the basinsconstitute a northern one. with the position of the river.lucar (7) somewhat ambiguous.

ami enciemie speeicsThis regionalization does not change when introducedspecies are excluded, because there is only one:Discojiiossus pie tus. However, when we consideredonly endemic amphibians (n = X). no regionalizationwas delected in the UPGMA analysis based on Jac-

eard"s index, whereas Baroni-Urbani and Buser'sshowed the existence of a strong biotie boundary be-tween basin 5 and the others, and a weak boundaryseparating the three south-eastern basins (7, H) andII) from the rest (Fig. 3j, Table 3). The pattern ob-tained using PAUP is similar to that obtained usingBaroni-Urbani and Buser's index, except that basin 9is associated with the south-eastern basins (Fig. 3k).The pattern supported by the consensus betweenPAUP and the other two trees is consistent with thatobtained using Baroni-Urbani and Buser's index (Fig.31),

The distribution pattern shared by fishes andamphibians

A combined components analysis of the tliree treesproduced tor all lishes and ihe three trees produced toral! ampliibians (Fig. 4a) resulted in pairs of basins withidentical associations for both taxa (basins 1-2, 3-6.8 9, 4 - 5 . and 10-11), with a frequency of occurrencetor interior nodes >93'Xi. The difference between thedistribution patterns of lishes and amphibians emerges

ilfOCIRAPHY :i 377

.3 "^

O

ca

&

O

c; c c c s = c

— r l —'

I 1 I

.0235

.5647

-

5400

5294

.0609

r-i

.6838

oc

5724

5254

,1955

.0915

.6297

,4212

0904

0434

3677

0655

oo oo o'ZJ oo

<:f r~- r- OS

d d ^ d

OS — •T '^ . o > . c ; 3 —r'l u-i O ' ^ oo OC oo r--,• ^ <-, VI >--; —; C — —

dd dd oddo

r-l r-l r-i ri

from the relationships among these pairs of basins. Inthe case of freshwater fishes, each pair of basins isrelated to the other basins llowing into the same sea(see. e.g. Fig. 2b). whereas in the case oi" ampliibiansthe pairs of basins are gi'ouped together according to alatitudinal pattern (I-'ig. 3b). The Jiiear basin (7). theonly basin that remains unpaired, is associated withthe Mediterranean basins based on fishes, but it isgrouped together with the northern basins ba.sed onamphibians.

The majority consensus tree between the patternstor indigenous lishes and amphibians (Fig. 4b) pro-(jiiees the same pairs of basins as above for all fishesand ail amphibians (F'ig. 4a). with a frequency ofoeeurrence of the pairs of basins even higher (>98'V;i).The consensus between the types ol' distribution (orendemic fishes and amphibians in the Iberian Penin-sula maintained only the pair of basins 10 and 11 andthe pair of basins 8 and 9, with also a trio of basins 2,3 and 6 with a frequency of occurrence>91'%. (F'ig.4c). The rest of the basins are associated with very lowfrequencies of oeeurrence.

Environmental factors related to the OBUs

The results of ihe discriminant analysis applied to theOBUs obtained for all iVeshwaier fishes showed thatthe environmental factor that best separated theCantabrian region (basins 1 and 2) from the otherregions (basins 3 1 I) is a higher run-off (RO) in theCantabrian region according t<:) the following discrimi-nant function:

Wilks" / = 0.0394;

p < 0.001. (I)

The enviroiuTienlal variables that best allow one todistinguish the Atlantic river basins (3, 6, 8 and 9)from the Mediterranean ones (4. 5, 7, 10 and II) arethe mean relative humidity at 07.00 for January (H),and the run-off (RO). which are both greater in theAtlantic basins, according lo ihe following functii^n:

y = -49.93-1-0.56//

p<0 .0! ,

Wilks" / =

(2)

Based on all amphibians, pluviometric irregularity(PI) is the variable best separating the northern riverbasins (1-7) wilh the lowest PI values from the basinsin the south (8 11) which display greater pluviometrieirregularity, according to tlie following function:

Y= -9.41 +0.32/^/

p<0 .01 .

Wilks' ;, = 0.3894:

(3)

378 21:4 (I'm)

43

79 Las-i100 1

1

77

h

S7

4-1

99

99 p

1 98 ] —99 1

10Q_i

1

fi?

10

48

53 ,

91 1—

100 1 •

Fig. 4. Consensus belwccii llie classificalioiis of Iberian riverbasins according to tresliwaler iisli iinci amphibian dislribu-lions using .laccard's index. Biironi-Urbani and Biiscr's index,and PAUP, lor all species (:0. indigenoLis species (b) andendemic species (c).

Discussion

Biogeographical regions based on all freshwaterfish species

The geographical distribution of freshwater fishes hasbeen used several times to divide the Iberian Peninsulainto biogeographical regions, although the methods usedlacked a probabilistic basis. The resulting patterns tendlo display two distinct configurations: ()ne showing afundamentaliy latitudinal division of regions (Lozani^1952, Alma(;a 1978, Doadrio 1988); and another indicat-ing the existence of biogeographical regions related to themain sea watersheds (Arevalo 1929. Hernando 1990.Hernando and Soriguer 1992).

Based on all Iberian freshwater fishes, our resultssupport the biogeographical regionalization related to ihethree major watersheds: the Cantabrian basins: theAtlantic basins; and the Mediterranean basins. TheCantabrian region is inhabited by fewer species than theother two regions, mainly due to the occurrence o\' fewerintroduced species (5 out of 14 in the Iberian Peninsula).Using only indigenous speeies. the biotic boundarybetween ihe Cantabrian basins and the rest of thePeninsula disappears. Therefore, the low number of fishspecies introduced into the Cantabrian basins is probablythe main reason for this biotie boundary. In the IberianPeninsula, fishes have been mainly introduced into reser-voirs, but on the Cantabrian rivers few reservoirs havebeen built due to the high availability of water in them,as evidenced by the high run-off of their basins, and thelow severity of floodings. Therefore, the low number offish species introduced into the Cantabrian basins mightbe ultimately eaused by the high run-off values (seediseriminant function I).

The second biotie boundary for all Iberian fishes isfound between the Atlantic and Mediterranean riverbasins and is best characterized by the difference in airhumidity in .lanuary at 07.00 which is greater in the

Atlantic basins, and by Lhe higher i un-off in the Atlanticrivers. The river Ebro (4) is more similar to the Atlanticrivers if one considers only its morning humidity inwinter, but when the average run-off of its basin is takeninto aceount, it elearly has Mediterranean characteristics.The humidity of lhe river basins in January at 07.00 maybe a surrogate for the size, form and fiow of lhe rivers.In large basins, with a high volume of water and broadvalleys, a ihermal inversion takes place at night duringthe winter, and this leads to the formation ol" morningmists in the low areas around the rivers. In contrast, inshorter and narrower rivers typiea! o[' the Mediterraneanarea, the thermal inversion affeets smaller areas, ihusreducing the average humidily of the river basins (Capel1981). Though it is not clear how this relates lo !ishphysiology, certain speeies are replaeed by others at thisbiotic boundary, and the environmental factors thatcoincide with ihis replacement are the morning humidilyin winter of the river basins, higher on lhe Allanticwatershed, and the average run-off, also higher in thebasins thai open inlo the Atlantic Ocean.

Biogeographical regions based on all amphibians

Previous studies have nol revealed the existence of anyclearly defined biogeographical regions in the IberianPeninsula using amphibians (Schall and Pianka 1977.Btisack and Hedges 1984). Busack and Jaksie (1982)attributed this to a hypothesized low ecological special-ization of amphibians in the Peninsula. However, weidentified a northern and a southern biogeographicalregion, separated by a weak biotic boundary. Fifteen of17 species o\' the southern region are present in thenorthern region, but only 15 out of 2? northern speeiesare present in the southern region. Thus, the boundarymay be considered '"semi-permeable" (Hernandez-Bermejo and Sainz-Ollero 1984), and we hypothesize thatdispersal is more difficult from north to south.

The environmental factor that best characterizes thisbiotic boundary is the greater interannua! pluviometrieirregularity (PI) ofthe southern region (see diseriminantfunction 3). The lower predietabilily of precipitation isa serious obstacle for amphibians that depend on seasonalrainfall for their reproductive cycle, and may preventsome species from inhabiting the southern region.

Biogeographical eonsensus between freshwaterfishes and amphibians

Reologically, the iwo patterns obtained using all speciesseem to represent mainly the spatial responses of fishesand amphibians to two different climatic processes; in theease of fishes the gradient of run-off creates a patternrelated to the major watersheds, and for amphibians thegradient o\' rainfall rregularity creates a north-southdivision.

However, eeoelimatie faetors alone cannot explain theconsenstis between both patterns. This consensus

FClXiRAI'lHY :i:4 379

can be seen in the formation of pait-s of river basins,which are identical for freshwater tish and atnphibianpatterns, using all species or using only indigenousspecies. The relationship between each pair of riverbasins might have a historical origin, because it is inagreement with the genesis of the Iberian river basinsin the Preglacial Age (see Introduction).

taxa: a) the formation of isolated land areas during theUpper Eocene-Lower Oligocene period (40- 30 Myr BP)in the case of amphibians, and b) the fonnation ofendorrheic basins during tlie Upper Oligocene-LowerMiocene period (30 23 Myr BP) in the case of tislies.

Acioiiwlcclfii'inculs We avL- grateful to J. Olivero and MiguelA. Ri;nd(Sii-Martos for llieir assistance vvitli the ligiires.

Biogeographical regions based on indigenous andendetiiic species

When only indigenous ftsh species are considered, aboundary is found between a Mediterranean biotic regionand a Cantabro-Atlantic region. This tnight have beenIhe result of different dispersal events during the LlpperMiocene transgression (Doadrio 1990). It is worth notingIhat the Pyrenees constitute an effective barrier to fishes,only permeable at its extremes. Freshwater fishes enteringthe Iberian Peninsula ftxim Hurope through the Mediter-ranean extreme of the Pyrenees likely colonized theMediterranean basitis. whereas those entering throughthe Atlantic side mostly colonized the Cantabro-Atlantierivers.

Using endemic fish species, our t-esults support thelatitudinal pattern. This may reflect the geographicalpattern of the Iberian Peninsula endorrheic basins duringthe Upper Oligocene-Lower Miocene period (30 23 MyrBP), when tnost endemic species arose (Doadrio 1990).During the Upper Oligocene the Pyrenees wet-e formed,constituting a barrier to fishes and enclosing severalendorrheic basins. This endorrheism mainly involvedwhat now corresponds to the basins of the rivers Diiero(3), Ebro (4), Tajo (6) and Jucar (7) (Lopez-Martinez1989). and stands out as one of the tnain factorsexplaining the appearance of many endemic species in theIberian river basins (Doadrio et al. 1991). The southernBctic-Riff Massif was a separated land area whicht-emained isolated, probably with its own endemic fauna,until the Messinian period in tiie Miocene, and eventuallycontributed to the formation of the current rivers Gua-diana (8), Guadalquivir (9), Segura (10) and Sur (11).

Using endetnic ampliibians. our biogeographical re-siionalizatioti reilects almost exactly the three separatedland areas of the Iberian Peninsula existing during theUpper Eocene-Lower Oligocene period (40-30 Myr liP)(Lopez-Martinez 1989). In this context, only Eupvoctusaspci- (Duges) would have originated in the north-easternland area, wliet-e the Pyrenees were rising (Caccone et al1994); Alyfcs dukhillcni would have originated in thesouthem Betic-Riff Massif (Arntzen and Garcia-Paris1995); the other 6 endemic amphibians, which occupymost of the current Iberian Peninsula, would havedifferentiated in the central land area.

The low degree of consensus between the biogeograph-ical patterns of endemic fishes and endemic amphibiansmay result from different processes influencing the two

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Appendix 1

Spceies

, Fresbwator Hsh species present in

Status

each

1

Iberian

-)

river basin.

3

1

4

= introdueed species: e =

River basins

5 6 7

endemic

K 9

species.

10 II

m phitu-rt'yzini nuircr siurio

.•ilo.su ahisdAh'sa fallax

iliu/iii ItiulwOncdihyitclnis niykissSaltiu/ salufSiilnin IrtiliaSalvcliitiis foiiiimilis

Aiiaci\y/iii.\ hispaniraBcirhtts hiiva^i-iBarhus coiaiictBar hits ^rctc/hiiBar hits ^tiirtionisBarhtts luui.\iBarhlts inciiilioiialis

+ +

FtXKiKAl'HY 1\A

Appendix I. (CorUiniied)

Species Suilus Rivtrr basins

K 9 10

Bailms iiiicrai't'plniliis e - 1 - 4 -Biirhiis silait'ii e 4 - 4 - 4 -<.\irii.\.\iii.\ auralus \ 4 - 4 - - I - 4 - 4 - + 4 - 4 - 4 - 4 -Cypiintis cm pic i 4 - 4 - 4 - 4 - 4 - 4 - 4 - 4 - 4 -ClioiklrosioiiHi polyk-pis e 4 - 4 - 4 - 4 - 4 - 4 - 4 -Cli/'iKlri'MoiiHi loxosUmtit 4- 4 - 4 - 4 -Gf'hia !-oNit i 4- 4 - 4 - 4 - 4 - 4 - 4 - 4 - 4 -Ilvrovypris pidiuinsi c 4-Lciuisciis caroliicnii c -I- -f 4- 4-Lciuisi'iis ivplkiliis 4- 4-Leiirisriis p\'i'ciHiicii!> e 4- -H 4- -|- 4-I'lloxiiius phoxiiuis 4- 4- 4- -t-

Riililiis h'ini!iiiii;ii e 4- 4 - 4 - 4 -Sdirditntts crylhroplulhilintis i 4- 4-•/•(•/((•(( liiica 4 - 4 - 4 - 4 - 4 - 4 - 4 -I'ropidiiphoxiiicllits alhmuoiilcs s 4- 4 - 4 - 4 -Ci'hili.s cethli-riini e 4- + 4-Cohiiis niiir/HTtina 4- 4 - 4 - 4 - 4 -.\iifnittliliciliis I'urhalulus 4- 4-Sihifus fiUiitis i 4-Aiin-iuni.i iiu'lus \ 4 - 4 - 4 - 4 -Aplumiiis iiicnis 4 - 4 - 4 - 4 - 4 - 4 -l-'iindtillis licU-focliliis i 4- 4-Wik'uciti hisjHiiiicu e -I- 4-(idinhiisiti liolhnHiki i 4 - 4 - 4 - 4 - 4 - 4 - 4 - 4 -Alhcriiui hoyeri 4 - 4 - 4 - 4 - 4 - 4 - 4-(kisicrosleus aculcanis 4 - 4 - 4 - 4 - 4 - 4 - 4 -

Micfopienis siiluioidcs i 4 - 4 - 4 - 4 - 4 - 4 - 4 - 4 - 4 - 4 -Blaniiiis flinialilis 4 - 4 - 4 - 4 - 4 - 4 -

Appendix 2. Amphibian species present in each Iberian river basin, i - inlrodiiced ^^pecies; e = endemic species.

Species Sialus River basins

1 2 3 4 5 6 7 8 9 10

Cliii<i:!ox.\a hisilaiiivii e 4- 4- 4- 4-

SaluDuiDilrti siiliiiuandm 4- 4-Flctirodcles wall IEitprocui.s cisper e 4-Triniriis marmoratus 4- 4-TrUiirus pyi;tii(tcii\- eTriiiiriis (ilpi'siiis 4- 4-Triiuriis hosioi e 4 - 4 -Triiiirus lu'lpciirus + +Disaisflo.^sits i-ah^diuii e 4 - 4 -Disciiiilosstis picliis iAlylc.s iihsU'lrivuiis -|- 4-Alvlcs dickhillcni eAlyli-s cisWinusii S 4-Pcliiliaics ciilnipcs 4- 4-I'clodyics imiiciuliis + 4 - 4 - 4 - -J- - 1 - 4 - 4 - 4 - 4 - 4 -Biijii biifo 4 - 4 - 4 - 4 - 4 - 4 - 4 - 4 - 4 - 4 - 4-Bufii calaiiiiUi 4- +Biijd rirtdisllvia m-hon-a 4- 4-Ilylci fiicridiiiiiiilisRii'iii pvrczi 4- 4-Riiiiit iliilnialina 4-Rmiu iiicrica s 4 - 4 -Riiiiii Icin/'itrariii 4- 4-

382

4-4-4-

4-

4-4-4-

4-

4-

+4-4-4-

4-

4--1-

4-4-4-4-

4-

4-4-

4-

++4-

+4-. f

4-

4-4-

4-

4--h

4-

•i-

4-4-

4-4-4-

4-4-4-

4-4-4-4-4-

4-

4-

4-4-4-4-4-

4-4--f

4-

4-

4-

4-4-

-H

4-4-4-4-

4-

+

+4-

4-

4-

4-

4-

4-4-4-

4-4-

+4-

+

4-4-

4-

4-

4-

4-

4-4-4-4-

4-4-4-

4-

4-

4-

4-

4-4-4-4-4-4-

4-4-