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EDAPHIC DIFFERENTIATION in LASTHENIA: a MODEL for STUDIES in EVOLUTIONARY ECOLOGY Author(S): Nishanta Rajakaruna Source: Madroño, Vol

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EDAPHIC DIFFERENTIATION in LASTHENIA: a MODEL for STUDIES in EVOLUTIONARY ECOLOGY Author(S): Nishanta Rajakaruna Source: Madroño, Vol EDAPHIC DIFFERENTIATION IN LASTHENIA: A MODEL FOR STUDIES IN EVOLUTIONARY ECOLOGY Author(s): Nishanta Rajakaruna Source: Madroño, Vol. 50, No. 1 (JANUARY-MARCH 2003), pp. 34-40 Published by: California Botanical Society Stable URL: http://www.jstor.org/stable/41425490 . Accessed: 18/10/2013 21:05 Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp . JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. California Botanical Society is collaborating with JSTOR to digitize, preserve and extend access to Madroño. http://www.jstor.org This content downloaded from 68.189.1.182 on Fri, 18 Oct 2013 21:05:19 PM All use subject to JSTOR Terms and Conditions Madroño,Vol. 50, No. 1, pp. 34-40, 2003 EDAPHIC DIFFERENTIATION IN LASTHENIA : A MODEL FOR STUDIES IN EVOLUTIONARY ECOLOGY Nishanta Rajakaruna Departmentof Biological Sciences, 371 Serra Mall, StanfordUniversity, Stanford,CA 94305-5020 nishanta@ stanford.edu Abstract Thecurrent knowledge of thegoldfield genus Lasthenia is discussed,emphasizing the possible role of edaphicfactors in thedivergence of thispredominantly Californian genus. Lasthenia species occupy a widearray of edaphic habitats ranging from serpentine outcrops to saltflats to guano deposits, thriving underspecific ion and osmotic conditions that are uninhabitable forthe vast majority ofspecies. Studies showthat ion accumulation and sequestration are commonstrategies used to counterosmotic effects, commonto mosthabitats harboring Lasthenia species. Inter- and intra-specific variation in toleranceto edaphicconditions has been observed in closely related yet reproductively isolated taxa, suggesting that edaphicfactors may have set the stage for the diversification ofthis genus. Lasthenia provides numerous opportunitiestoexamine the link between adaptation, reproductive isolation and speciation. KeyWords: adaptive divergence; California flora; edaphic races; Lasthenia ; salinity tolerance; serpentine. Accordingto Lindleyand Moore (1876), "they 1989; Desrochersand Böhm 1993) to assess bio- ( Lasthenia) growin wet places,and appearto be systematicrelationships and determinetrends of uninterestingweeds." Theydo "growin wetplac- biochemicalevolution in thegenus. Electrophoretic es," but studiesof Lasthenia(Heliantheae: Astera- workfollowed to establishevolutionary relation- ceae) overthe last severaldecades have shownthe shipsamong closely related taxa withinthe genus genus to consistof anythingbut "uninteresting(Crawford et al. 1985;Crawford and Ornduff 1989) weeds." Commonlyknown as goldfields,plants of as well as withinthe highly variable L. californica Lastheniaoccupy large areas of the Californian land- sensuOrnduff (Desrochers and Böhm 1993).More scape,casting spectacular carpets of brightly-colored recently, comprehensive phylogenetic studies goldenyellow flowers in earlyspring. A recentmo- (Chan et al. 2001, 2002; Desrochersand Dodge lecularphylogenetic study recognized 21 speciesand 2003) havecontributed to ourunderstanding ofpat- subspeciesbelonging to sevensections (Chan et al. ternsof divergencein Lasthenia.The studyby 2001). All butone speciesare endemicto theCali- Chan et al. (2001) agreeswith observations made fornianFloristic Province. Lasthenia kunthii (Less.) earlier(Ornduff 1966, 1976) thatdivergence pat- Hook & Arn.,the only member of thegenus found ternsin thegenus conform to expectationsof cat- outsidewestern North America, is endemicto vernal astrophicselection and saltationaldiversification pools and wetlandsin centralChile (Ornduff 1966). (Lewis 1962). Membersof thegenus have wideedaphic tolerance Existingknowledge suggests that Lasthenia can andare found in habitatssuch as coastalbluffs, gua- providea modelsystem for studies in evolutionary no deposits,vernal pools, salt and alkalineflats, ser- ecology,specifically, in understandingthe role of pentineoutcrops, deserts, grasslands, and open edaphicfactors in differentiation.Species of Las- woodlands(Ornduff 1966, 1993). Membersof L. theniahave successfullycolonized diverse habitats californica sensu Ornduff (Ornduff 1993) have the withinCalifornia, including those that exclude the widestedaphic tolerance within the genus, with pop- vast majorityof species (Ornduff1966, 1993; ulationsspanning all but guano habitats.Keck Kingsburyet al. 1976;Vasey 1985; Rajakaruna and (1959) statedthat L. californica sensu Ornduff [then Böhm 1999; Noe and Zedier 2000; Parsonsand Baeria chrysostoma(Fischer & C. Meyer) E. Whelchel2000). Further,in some sections(e.g., Greene]was themost abundant composite in Cali- sect.Hologymne [Battling] A. Grayand sect.Orn- fornia.Other Lasthenia taxa have ratherrestricted duffia R. Chan)all speciesoccupy the same edaphic distributions,with seven taxa now listedin Califor- habitatyet generally do notgrow intermixed (Orn- nia NativePlant Society's "Inventory of Rare and duff1966), whilein others(e.g., sect. Ptilomeris EndangeredPlants of California"(CNPS 2001). [Nutt.]Ornduff), species occupy contrasting edaph- Following Ornduff's extensive monograph ic habitats(Crawford et al. 1985; Vasey 1985). (1966), Lastheniahas receivedconsiderable atten- Thusit is plausibleto hypothesizethat edaphic fac- tion.Early studies examined inter- and intra-specif- tors may have playedan importantrole in thedi- ic variationin flavonoidchemistry (Saleh and versificationof thegenus. In thispaper, taxa in sev- Böhm1971; Ornduffet al. 1973,1974; Böhm et al. eral sectionsare discussedwith emphasis on the This content downloaded from 68.189.1.182 on Fri, 18 Oct 2013 21:05:19 PM All use subject to JSTOR Terms and Conditions 2003] RAJAKARUNA:EDAPHIC DIFFERENTIATION IN LASTHENIA 35 possiblerole of edaphicfactors in theirdiversifi- and L. gracilis,suggesting that racial features may cation.Since manyendemic species of flowering have evolved secondarilyin responseto edaphic plantsin westernNorth America are edaphicen- factors(Rajakaruna et al. in pressa). demics(Kruckeberg 1969, 1986),studies of a mod- Recenteco-physiological studies (Rajakaruna et el genussuch as Lastheniacould providegeneral al. 2003) indicatethat race A plantsfrom both L. insightsinto speciation in a varietyof othersuch californicasubsp. californicaand L. gracilisare edaphicallydiverse genera. clearlymore tolerant of potentially-toxicNa+, sug- gestingthat physiological traits responsible for Na+ Lastheniasect. Amphiachaenia (DC.) R. Chan: uptakeand sequestrationmay have evolvedinde- L. californicasensu Ornduff pendentlyin populationsbelonging to the two closelyrelated taxa. Schat et al. (1996) provideone Lastheniasect. Amphiachaenia, formerly known of thebest examples of parallelgenotypic changes as L. sect. Baeria (Fisch. & Mey.) Ornduff,was in toleranceto an edaphicextreme within a plant recentlyrevised (Chan 2001) to includefour spe- species.Their studies have shownthat loci confer- cies,L. californicaDC. ex Lindl,[consists of subsp. ringcopper tolerance have evolvedindependently californica,subsp. macrantha (A. Gray)R. Chan, in geographicallyisolated populations of Silene vul- and subsp.bakeri (J. T. Howell)R. Chan],L. grac- garisGarcke (Caryophyllaceae). In Plantago(Plan- ilis (DC.) Greene,L. ornduffiiR. Chan,and L. lep- taginaceae),the vacuolar Na+/H+ antiporter activi- talea (A. Gray)Ornduff. Lasthenia californica and ty,thought to confertolerance to Na+ (Apse et al. L. graciliswere previously recognized as L. cali- 1999) is onlypresent in thesalt-tolerant P. mariti- fornicasensu Ornduff (1993). The twocryptic taxa ma L. butnot in theglycophytic P. mediaL. (Staal foundwithin L. californicasensu Ornduff, L. cali- et al. 1991).This difference is thoughtto be crucial fornica subsp. californicaand L. gracilis,have in theecological divergence of thesetwo species. been the subjectof intenseecological and evolu- Sodiumis clearlyan importantelement in manyof tionarystudies (Rajakaruna and Böhm 1999; Chan the habitatswhere race A is found.Whether the et al. 2002; Desrochersand Dodge 2003). presence/absenceor levelof expressionof theNa+/ A biosystematicstudy (Desrochers and Böhm H+ antiportergene is responsiblefor the differences 1995) firstsuggested the existenceof two geo- in uptakeand accumulationin race A plantsis an graphicalraces within L californicasensu Ornduff area worthyof investigation.Ion uptakestudies based on flavonoidpigments (Böhm et al. 1989; conductedon thetwo races from Jasper Ridge also Desrochersand Böhm 1993), allozymebanding indicatethat race A is moretolerant of highexter- patterns,and cypsela characteristics.A detailed nal Mg2+and low Ca2+/Mg2+ratios (Rajakaruna et ecologicalstudy (Rajakaruna and Böhm 1999) sug- al. 2003). Bothof thesetraits have adaptivesignif- gestedthat the two races describedby Desrochers icance in magnesium-richserpentine and coastal and Böhm(1995) occurin distinctsets of habitats habitatswhere race A is found.The traitsthat have thatcan be classifiedon thebasis of ionic stresses been studied(Na+, Mg2+,Ca2+/Mg2+) are common and wateravailability. Race A plantspredominate featuresin mosthabitats where race A is found, in habitatssubject to ionicstress. Although the soils thus,it is temptingto hypothesizethat differential in theseenvironments are ionicallyharsh,
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