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Marine Phytogeography of the Juan Fernandez Archipelago PDF

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PacificScience(1992), vol. 46, no. 4:438-452 © 1992byUniversityofHawaiiPress. AlI rightsreserved Marine Phytogeography ofthe Juan Fernandez Archipelago: A New Assessment! B. SANTELICES2 ABSTRACT: Anewassessmentofthegeographicaffinitiesofthe marinealgae of the Juan Fernandez Archipelago indicates a flora with a small number of species and very high endemism (about 30%) as compared to other oceanic islandsofsimilarage,size,origin,andabioticconditions.Thefloraalsocontains many widely distributed species (45%) and a small group of species with circumpolar-subantarctic affinities (about 13.5% of the flora). The potential algal species sources for this flora seem to be distant localities in the southern Pacific, including the southern tip ofSouth America, southern Australia, New Zealand, and several subantarctic islands. Considering effective dispersal dis tancesofmarinebenthicalgae,theJuanFernandezArchipelagoappearsasmore isolated than Easter Island, which previously was supposed to be the most isolated point in the Pacific basin. An analysis of the endemic components suggests that there has been speciation but no radiation in these islands. Some species originatingin thearchipelago might have migrated across the Pacificto continentalSouthAmerica, perhapsvia EINino/SouthernOscillation (ENSO). BASED ON THE SEAWEEDS collected by the by the abundance of subtropical elements, Swedish expedition to Juan Fernandez and which were common in Peru and northern Easter Island, Levring (1941) and Skottsberg Chile. Therefore, Skottsberg (l943a) con (1943a,b) made what amounts to be the only cludedthattheislandflorawasmorelikethat phytogeographic characterization ofthe ma ofPeru and northern Chile than that ofthe rine algae of the Juan Fernandez Archipel continental flora farther south. ago. They distinguished eightfloristic groups EventhoughSkottsberg(1943a)recognized among the nearly 100 seaweed species re that the West Wind Drift and the Chile-Peru ported from the islands. Endemic elements Current (Humboldt Current) was the route weremostabundantlyrepresentedintheflora probably followed by a large number of (32%), followed by subcosmopolitan (20%), species, he considered that the abundance and subtropical species widely distributed in of subtropical elements in this insular flora warm waters (14%). They also noticed the was greater than what could be attributed to presence of a small (10%) Australasian temperatureconditionsalone. He(Skottsberg Neozelandian element that did not belong to 1943a)furthersuggestedthathistoricalcauses the circumpolar group known also from had been influential. Considering that the southern Chile and Magellania, and a few fossillandfloraofsouthernChileandAntarc (about 5% ofthe flora) subantarctic species. tica showed that during the Tertiary the cli Both Levringand Skottsbergwere impressed mate was considerably warmer than now, with the absence in the islands of the large Skottsberg supported the idea that Juan kelpsandbullkelpsthataresocommoninthe Fernandez was formerly connected with southern part ofPacific South America and South America. He disregarded a neovo1ca nic, oceanic origin of the Juan Fernandez Islands because seemingly it did not allow 1Manuscriptaccepted 18January 1992. enough time for the evolution of strongly 2Departamento de Ecologia, Faculated de Ciencias marked endemic forms and because a major Biol6gicas, P. Universidad Cat61ica de Chile, Casilla 114-D,Santiago,Chile. ity of the endemic species belong to genera 438 MarinePhytogeographyoftheJuan FernandezArchipelago-SANTELICES 439 regarded by Skottsberg (1943a) as having 1973).AccordingtoArana(1987)thismodifi tropical and subtropical affinities. cationcouldhavebeenrelatedtotheaperiodic In the 50 yr following Skottsberg's pro ElNino/SouthernOscillation(ENSO), which posal, several expeditions have visited the wasexceptionally strongthat year. islands, increasing the geological, oceano Marine floristic studies have added new graphic, and floristic knowledge ofthe archi perspectives to the biogeographic affinities pelago. It is now accepted that the Juan of the flora. Even though new collections Fernandez Islands are entirely volcanic in (Etcheverry 1960, Earle 1969, Hawkes and origin and apparently have never been con Johnson 1981)haveadded few records to the nected to the mainland (Stuessy et al. 1984, islands, floristicstudieselsewhere have modi Gonzalez-Ferran 1987). The Nazca Plate is fied the known geographic distribution of movingfrom westto eastand apparentlynew several of the species found in the Juan islands are produced as the plate moves east Fernandez Archipelago. Several of the sub wardovera"hotspot."Theabsoluteagesare tropical elements considered as most impor estimated at 3.8 to 4.2 million years (My) tant by Skottsberg(1943a) are now regarded for Masatierra Island and I to 2.4 My for as widespread bothin temperateand tropical MasafueraIsland.Theknownspreadingrates waters. Some of these taxa also have been forthemarginoftheNazcaPlate(8.8cmyr-1 found in southern South America (see re ) agree well with the observed distance sepa view by Ramirez and Santelices 1991), in rating Masatierra and Masafuera (150 km) creasing the floristic affinities between the and their known ages. Therefore, present Juan FernandezArchipelagoand localitiesin geological data do not support Skottsberg's southern South America, New Zealand, and (1943a) proposal ofa land connection ofthe Australia. Juan FernandezIslands with SouthAmerica, Ontheotherhand, newdataonthefloraof and the argument cannot be used to explain EasterIsland, anotheroceanicisland ofsimi thegeographicaffinitiesofthemarineflora of lar age, size, origin, and approximately simi the islandswith that ofthe continent. lar abiotic conditions as those in the Juan Oceanographicstudieshaveconcludedthat FernandezArchipelago, areinconsistentwith the surface waters surrounding the Juan ourpresentunderstandingoftherelationship FernandezIslandsaremainlyofsubantarctic among isolation, species richness, and en origin. The most superficial layer, and down demism,butareusefulforcomparison.Easter to 200 m depth, corresponds to the subant Islandhasbeenfound tohaveabout 1/3more arctic water mass, with temperature ranges species but only about halfthe percentage of from 10to 19°Candsalinitiesbetween34.00/00 endemic elements as the flora of the Juan and 34.2%0. Furthermore, the ocean circula Fernandez Archipelago (Santelices and Ab tion around the islands, between 75° Wand bott1987).Alloftheseislandarevolcanicand 78° W, is under the influence of the strong ofabout thesameage(3.0 My, EasterIsland, northward flow of the Humboldt Oceanic Newman and Foster 1983) and the same size Branch, also called the Chile-Peru Current (Masatierra = 93 km2, Masafuera = 60km2, (Figure I). Between 78° Wand 80° W, there Easter Island = 93 km2 All are hilly and ). is a strong southward flow that may corre relatively barren, with coastlines character spond to the Peru Oceanic Countercurrent; ized by cliffs; few, small beaches; and few beyond 81°W, acurrentflowing to the north sheltered bays(Stuessyetal. 1984, Gonzalez withlowvelocityandsmallvolume transport Ferran 1987). Winds are strong and change has been identified (Arana 1987). directions seasonally and daily (Hajek and Onatleastoneoccasionthisgeneralsubant Espinoza 1987). Thus, there seem to be no arctic influence was found to be altered: the obviousdifferencesin theabovedeterminants most superficial 50 m around the islands was ofspeciesrichnessandendemismbetweenthe found in April 1973 to beoccupied bywaters islands in the Juan Fernandez Archipelago of the subtropical water mass, with higher and EasterIsland. temperaturesand salinities (Silvaand Sievers The most important difference between 440 PACIFICSCIENCE, Volume46, October 1992 110 100 90 80 70 0..(.'....0..".". •,,-. "\A'\ \ i\ ~1~ "0.~('~ " \0;1 '~ ~O, :'\ C~" f' 20 (~' o,. ~.~ ~;':)\ 0 20 ~ 1t ~ ; ~ '\0/\~ ~\ ~~lJ! t 0 q Easter Island 30 ~ '~J 30 t FernanCf(.7./~ ! / J. /1. --.::, / 40 • " ~ 40 West Wind Drift 50 50 110 100 90 FIGURE I. Pattern ofoceanographiccirculation around the Juan Fernandez Archipelago (redrawn from Arana 1987). Easter Island and the islands in the Juan (Briggs1974)tobeconsiderablylessthanthat FernandezArchipelagoseemstobethedegree ofEasterIsland.However,thehighendemism ofisolation.EasterIslandisrecognizedasone and the small number ofspecies exhibited by ofthemostisolatedpointsinthePacific,lying the flora ofthe Juan Fernandez Archipelago 3250kmawayfromtheSouthAmericancoast as compared to Easter Island are not consis and 2315 km away from the nearest land to tentwith their respectivedegrees ofisolation. the west (Ducie Atoll). The Juan Fernandez Thedegree ofendemismand the speciesrich Archipelago is only 600 km away from the nessofJuan Fernandezmightbeexplainedby South American continent. Therefore, the dispersion distances and biological isolation degree of isolation of the Juan Fernandez severalordersofmagnitudegreaterthanthose Archipelagohastraditionallybeenconsidered in effect on Easter Island. This assumption is TABLE I EXAMPLESOFPATTERNSOFGEOGRAPHICDISTRIBUTIONSOFBENTHICALGAEREpORTEDFROMTHEJUANFERNANDEZARCHIPELAGOANDEASTERISLAND JUANFERNANDEZ EASTERISLAND REPRESENTATIVELOCALITIESELSEWHERE Indo-PacificDistribution Cladophorasocialis + Tahiti,Polynesia,Malaysia Cladophoropsisherpestica + Vietnam,Japan,southernAustralia Ectocarpusarabicus + + IndianOcean,India,Malaysia Feldmanniaindica + Indo-Pacific,Indonesia,Philippines,HawaiianIslands Sphacelariatailensis + Tahiti Dictyotacrenulata + CentralPacific,Philippines,HawaiianIslands Dictyopterisrepens + WesternPacific,Japan,Philippines,MarianaIslands Lilhophyl/umsamoense + Tahiti,Samoa,Hawaii Ceramiumcruciatum + Indo-Pacific Polysiphoniasavatieri + TropicalPacific,Hawaii,Malaysia,Japan Circumpolar-SubantarcticDistribution Scytothamnusaustralis + CentralsouthernChile,Australia,Tasmania,NewZealand Splachnidiumrugulosum + Tasmania,NewZealand,SouthAfrica Glossophorakunthii + SouthernPerutoCapeHorn,ChathamIslands,NewZealand Acrochaetiumcatenulatum + + CentraltosouthernChile,CampbellIslands,NewZealand Gymnogongrusfurcel/atus + CentralPerutosouthernChile,NewZealand,Australia Rhodymeniaaustralis + Australia Rhodymeniacuneifolia + ChiloetoMagellanStraitinsouthernChile,FalklandIslands, southernArgentina Phycodrisguercifolia + CentraltosouthernChile,FalklandIslands,KerguelenIslands, SouthGeorgiaIslands,andCampbellIslands Schizoserisgriffithsia + ConcepciontoMagellanStrait,FalklandIslands,NewZealand, AucklandIslands,ChathamIslands,CampbellIslands Polysiphoniaabscissa + CoquimbotoMagellanStrait,severalsubantarcticislands EndemicSpecies Phaeophilapacifica Ectocarpuschnoosporae Phaeophilaramosa Sargassumskottsbergii Codiumfernandezianum Erythrocladialaurenciae Codiumcerebriformis Erythrocladiarepans Ectocarpusminutissimus Acrochaetiumdiscoideum Distromiumskollsbergii Acrochaetiumralfsiae Padinafernandeziana Galaxaurapaschalis Leptophyl/umfernandezianum Botryocladiaskollsbergii Chondriel/apusilla Ceramiumskollsbergii Hypoglossumparvulum Laurenciaclavata 442 PACIFICSCIENCE,Volume46,October 1992 consistent with the notion that the majority of the floristic elements present in the is lands ofthe archipelago originate in Fuegia MagellaniaandtheAustralasianarea,involv ing dispersal distances of 3000 (Fuegia) to 12,000 km (Australia-New Zealand). This relation was perceived by Skottsberg (1943a) but was overlooked by him because he supported a land connection between Juan Fernimdezandthecontinent.Theabundance oftropicalelementsinthearchipelago,andits generalfloristic affinity with Peru and north ern Chile, can be explained in two ways: (1) Some species ofwidedistributioncould have arrived independently in the islands and the continent; (2) Species restricted to Peru northern Chile and Juan Fernandez could have been transported across the eastern Pacificby aperiodicENSOevents. Inthisstudytheseideasweretestedandthe geographic affinities of the marine flora of JuanFernandezwerereassessed. Forcompar ative purposes, the marine flora of Easter Island has beenincludedin the analysis. ! + I + I I I ~ .~ METHODS This study is restricted to the benthic ~ Chlorophyta, Phaeophyta, and Rhodophyta recorded in the Juan Fernandez Archipelago andonEasterIsland. Dataonthegeographic distribution of the species were taken from the literature, most of which has been sum marized by Santelices (1987), Santelices and Abbott (1987), and Ramirez and Santelices (1991) for the areas in question. Data were initially recorded for 110 taxa in Juan Fernandez and 165 taxa on Easter Island. Taxonomic uncertainties or lack ofinforma tion on geographic distribution elsewhere re duced thenumberofspeciesconsidered to 89 inJuan Fernandez and 105 on Easter Island. Our samples represent 80% and 65% ofthe respective total number oftaxa reported for theseislands, and theproportionofendemics to total number of species is only slightly differentfrom thevaluescorrespondingtothe totalflora. Thespeciesweregroupedaccordingtotheir patterns of geographic distribution in the MarinePhytogeographyoftheJuan FernandezArchipelago-SANTELICES 443 Pacific basin and elsewhere. Table I shows a ure2).Thenumericallymostimportantgroup few outstanding examples of each group of includesspecieswithwidedistribution, which speciesdistinguished in each location. Abun compose 45-50% of the flora of these is dance of algae groups among locations was lands. Endemic elements are important in compared bychi-squaretests. bothplaces,buttheyareproportionallymore important in the Juan Fernandez Archipel ago. A small group ofspecies that extends to the Pacific coast of South America is also RESULTS moreimportantintheJuan FernandezArchi Fivegroupsofspeciescan bedistinguished pelago. The two remaininggroups, taxa with when the bulk ofalgal taxa reported for the Indo-Pacificaffinitiesandspecieswithcircum Juan Fernandez Archipelago and Easter Is polar-subantarctic affinities, also are pres land is grouped by geographic affinities (Fig- ent in both areas but their representation in EASTER ISLAND J. FERNANDEZ 60 40 20 0 20 4.0 60 WIDELY 49.5 44.9 DISTRIBUTED INDO- PACIFIC 28.6 2.2 J13' CIRCUMPOLAR 3.8 SUBANTARTIC ENDEMIC 13.3 29.2 0 ISLES- PACIFIC 1.9 67 SOUTH AMERICA . OTHER 2.9 3.5 FIGURE2. Relativeimportanceofseveralfloristicgroupswithdifferentpatternsofdistributioninthemarineflora ofEasterIslandandtheJuanFernandezArchipelago. 100 160 180 160 140 80 \1 100 120 140 160 180 160 140 120 100 80 CURRENT SySTEM ~ FIGURE3. Distributionof30speciesofbenthicalgaewithaffinitiesintheIndo-Pacific.Numberincirclesrepresents themeannumberofspeciespresent. 100 120 140 160 180 160 140 120 100 80 60 ~ ~o : ! .' '.~,\\'" ;'~.. ..... ~. ~., ~::.sa~~~·J., :..,:~. ":~":~;':i"~ "'.""- . o .. ,.(. '. •• ~ ~ '. '. ~:.!).•.-....;".. , ":I/e"'.P.'• 1 ••:, ...t.. ...\ ....3,../.. 20 '. ,'. ":0<.~~ ,.;,:'.-...:.: ..'.-"::...",... ~ '..:", t....:~""...~~:;1'..•....•. 13\ "',\""\...\3'.-,-/ '•••O' (. '. ~ I \\ .~ EASTER ISLAND: 112\6 / Q -" ~~. _ A\ J FE~NAN~E7' I _ '--....."--"-" .. \ / 6 \.~) -_ .. ,,/ I 8 ~. -"-...e.- " ,CH~THAM / / :\~ ___.. _".-." -'.1 .................""........... 5· (<- -._..__.•- ...../.",,... .",.. _.._...._.-..._..._ .::..-::--/,' / KERGU_E0L'E-N""__.o. '"'-- ....... ....... ";_'C.A.M -BE"L-L' .. _00 /.~"_"_.. _... . /'.'. '8..........".........._"'.. \J-J ....... .'--'" -" --...- .. -_.. _..,../' ~ -.~ -......."-"-" .-.'".'- -..'.".,/... /'./:";.. '.-.. ............. _ ... - __.. _.. --e.--... _._/' ~../' I 2 f.' ,,~ /' ';':. _ ,,-?/' / .' - ... ... 100 120 160 140 120 100 80 60 _._.._.._ ...... Current system FIGURE4. Patternofdistributionof12speciesofbenthicalgaewithsubantarctic-circumpolardistribution.Number incirclesrepresentsthemeannumberofspeciespresent. 446 PACIFICSCIENCE,Volume46,October 1992 each place is very different. The Indo-Pacific in distant places in the southern hemisphere, elementsare very abundanton EasterIsland; a pattern ofdistribution that is closely asso those withcircumpolar-subantarcticaffinities ciated with the West Wind Drift and the are more abundant in Juan Fernandez. Al Chile-Peru Current systems (Figure 4). Eight though the endemic elements usually are re ofthesespeciesarefound bothat localitiesin gardedashighlycharacteristicofagivenarea, southernmost Chile and in distant places in these last two groups, with differential repre the southern Pacific Ocean, including south sentation and strong asymmetry among the ern Australia, New Zealand, Campbell Is islands, may better reflect the geographic lands, and ChathamIslands. Four other spe affinitiesofthelocalfloras. Theywilltherefore cies occur in these distant localities in the beconsidered first. Pacific, but have not been found in southern South America. Because all of these species seem to have a similar dispersal route and Species with Indo-Pacific Affinities only a few are restricted to localities in Aus This group of species is the second most tralasia or the subantarctic, it is unnecessary numerousonEasterIsland(Figure2),amount to distinguishsubgroupssuchassubantarctic ing to 28.6% of the flora in the sample andAustralasian-NeozelandianasSkottsberg considered for this study. This group com (1943a) did. Moreover, only two or three of bines species previously characterized by thesespeciesarealso found in northern Chile Santelices and Abbott (1987) as having a andPeru,showingahighersimilaritybetween western Pacific pattern ofdistribution and a JuanFernandezandsouthernChilethanwith few taxa previously included among those morenorthernlocalities. Infact, thepresence widely distributed in warm waters. Some of of these species in northern Chile and Peru the species in the group (Figure 3) are re doesnotrequireacirculationsystembetween stricted to a few islands in Polynesia; others the islands and the continent because they are present also in Malaysia, northeastern couldhavereachedtherefromsouthernlocal Australia, Vietnam, Japan, and the Philip itiesbymeansofcurrentsalongthecontinen pines,andafewreachnorthernNewZealand. tal margins. Thirty of these species are found on Easter Threespecieswithcircumpolar-subantarctic Island, butonlytwo ofthem arefound in the affinitiesoccuronEasterIsland.Interestingly, Juan Fernandez Archipelago. Furthermore, two of them (Codium pocockiae and Dasya one of these two taxa (Laurencia clavata) villosa) are absent from the Juan Fernandez couldhavearrivedinthearchipelago through Archipelago, illustrating the very limited ex the West Wind Drift and the Chile-Peru change between Easter Island and this archi Current system, because it is also found in pelago. southernAustralia.Thedifferenceinthenum ber of species of this group represented on Species with Wide Distribution Easter Island and in the Juan Fernandez Archipelago emphasizes the influence of the In both insular floras discussed here the Humboldt system on Juan Fernandez, which numerically most important group ofspecies with its cold temperature and northern flow are taxa with wide geographic distribution acts as a powerful barrier to dispersal for a (Figure 5A). About 17 of these species are majority ofspeciesfrom EasterIsland. subcosmopolitan, including Juan Fernandez andEasterIsland,andtheirgeographicdistri butions do not provide additional clues on Species with a Circumpolar-Subantarctic their geographic origins and dispersal. An Pattern ofDistribution other 33 species are found on Easter Island TheJuan Fernandez Archipelago contains butnot inJuan Fernandez (Figure 5B). They agroupof12specieswithgeographicaffinities exhibit wide distribution, in tropical and FIGURE5. Regional affinitiesofinsularspeciesofbenthicalgaewidelydistributed inwarmand temperatewaters inthePacificbasinandelsewhere.Numberincirclesrepresentsthemeannumberofspeciespresent. 120 140 160 180 160 140 120 20 WIDELY DISTRIBUTED IN . 20 ". WARM AND TEMPERATE WATERS .. 0 ....:.. :·E>:· I 3 .. . '\ ~ , ....... '0·:\};.i.. 5, t'·-'.:":,.. ~t~. .' " ..' ® 6J 20 I i Easterisland ~ ~ Fernan at J1, I 6 I , 60 12il 140 160 180 160 140 120 100 80 60 DISTRIBUTED IN ',. '" SUBTROPICAL i:. 1.:,-. ... ··e······ .•... \:, ~\,'.,; " ..... .;.(.~~j:(.:'. , Ilio::-.. '" ..... ® 2I 20 ~I Easter island ., 2 .~ J Ferna ez t' i 40 ti rjJ. I, 120 140 160 1 0 160 140 120 100 80 DISTRIBUTED IN ....'. 20 TEMPERATE WATERS 0 ,. ......: ~" ':" ...~: .~ '. ;.:'1:·:;;~.~" :.'";:... ~1..: .. ..... ., ® 20 ;.' Easter island ( .~p .~ ... 68 120 140 160 180 160 140 120 100 80 60

Description:
Tropical Pacific, Hawaii, Malaysia, Japan. Circumpolar-Subantarctic Distribution. Scytothamnus australis. Splachnidium rugulosum. Glossophora
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