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Potential Ecosystem-Level Effects of Genetic Variation Among Populations of Metrosideros Polymorpha from a Soil Fertility Gradient in Hawaii

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Potential Ecosystem-Level Effects of Genetic Variation Among Populations of Metrosideros Polymorpha from a Soil Fertility Gradient in Hawaii Oecologia (2001) 126:266–275 DOI 10.1007/s004420000523 Kathleen K. Treseder · Peter M. Vitousek Potential ecosystem-level effects of genetic variation among populations of Metrosideros polymorpha from a soil fertility gradient in Hawaii Received: 7 December 1999 / Accepted: 22 August 2000 / Published online: 24 October 2000 © Springer-Verlag 2000 Abstract This study assessed intrinsic differences in tis- Keywords Allozymes · Common garden · Genetic sue quality and growth rate among populations of Met- variation · Nutrient cycling · Tissue quality rosideros polymorpha native to sites with a range of soil fertilities. We collected seedlings from three Hawaiian mesic forests that were either phosphorus-limited, nitro- Introduction gen-limited, or relatively fertile. These individuals were grown in a common garden under a factorial high/low, In ecosystem science, the composition (i.e. genotypes) of N/P fertilization regime for 1.5 years and then harvested plant communities has long been recognized as an im- to determine genetic divergence; aboveground growth portant factor in nutrient dynamics (Jenny 1941, 1980), rate; and lignin, N, and P concentrations in leaves and much recent attention has been focused on the roles and roots. Allozyme analyses indicated that the three of different species or functional groups (e.g. Wedin and groups had genetically diverged to some degree (gene- Tilman 1990; Hobbie 1992; Binkley 1995; Chapin et al. tic distance=0.036–0.053 among populations). Relative 1997; Steltzer and Bowman 1998). However, intrinsic growth rate did not differ significantly among the popu- differences in plant traits that could affect ecosystem lations. Senescent leaves from the fertile-site population processes (such as morphology, physiology, and alloca- had the highest N concentrations (due to low N resorp- tion) also occur among populations within species (e.g. tion) and had lower lignin concentrations than plants Clausen et al. 1940; Gauhl 1976; Chapin and Chapin from the N-limited site. Across treatments, P concentra- 1981; Eriksen and Nordal 1989; Norton et al. 1995; tions in senescent leaves were highest in plants from the Mansfield et al. 1999). Few studies have assessed the fertile and P-limited site. Root tissue quality did not gen- consequences of this intraspecific variation on ecosystem erally differ significantly among populations. Since de- function; one notable exception is the finding by Holland composition rate of senescent leaves in this system is re- and colleagues (1992) that populations of grasses with lated positively to N concentration and negatively to lig- different intrinsic grazing tolerances have distinct effects nin concentration, senescent leaves from the fertile-site on productivity and nitrogen mineralization. population may have a genetic tendency toward faster We examined intrinsic differences in plant popula- decay than the others. The intrinsic qualities of the three tions from sites that varied in soil nutrient availability, populations may provide positive feedbacks on nutrient and focused on traits that should feed back to affect nu- cycling at each site–nutrient availability may be raised to trient cycling. Plant traits that vary among species and some degree at the fertile site, and reduced at the N- or could potentially vary among populations include nitro- P-limited sites. Our results suggest that even a small de- gen (N), phosphorus (P), and lignin concentrations of gree of genetic differentiation among groups can influ- plant tissue; nutrient resorption upon tissue senescence; ence traits related to nutrient cycling. relative growth rates; and allocation strategies (e.g. Grime 1977, 1979; Chapin 1980; Coley et al. 1985; Chapin et al. 1986, 1990, 1993; Chapin 1991). Each of K.K. Treseder (✉) · P.M. Vitousek these characteristics is known to affect nutrient dynamics Department of Biological Sciences, Stanford University, through influences on tissue decomposition and/or eco- Stanford, CA 94305, USA e-mail: [email protected] system productivity. Fax: +1-909-7874625 We focus on Metrosideros polymorpha (Myrtaceae), Present address: an evergreen tree that dominates most rain forests in Ha- K.K. Treseder, Center for Conservation Biology, waii (Kitayama and Mueller-Dombois 1995). Like many University of California, Riverside, CA 92521, USA Hawaiian plants, Metrosideros occupies a wide range of 267 habitats within the archipelago, some with very different oped for allozyme analysis of M. polymorpha (Aradhya et al. selection pressures (Carlquist 1980). In addition, gene 1991, 1993). Briefly, about 25 mg of fresh leaf tissue was ground in 0.1 ml extraction buffer (Aradhya 1992) and absorbed onto fil- flow among the islands is sufficiently restricted to allow ter paper wicks. Wicks were loaded into 12% starch gels, and elec- differentiation to occur, and many plants and animals, in- trophoresis was conducted at 4°C at 300 V and 40 mA for 6 h. The cluding Metrosideros, have evolved island-endemic vari- gels and running buffer were prepared with a histidine-citrate pH eties, subspecies, or species (Carlquist 1980; Kitayama 6.5 solution. Gels were stained for eight enzyme systems (Ta- ble 2). Genotypes were inferred from allozyme phenotypes. and Mueller-Dombois 1995). Genetic differentiation among populations was assessed by de- To examine intrinsic differences among M. polymor- termining the genetic distance between pairs of populations. Allele pha populations from areas with a range of soil fertili- frequencies were calculated for each population, then the genetic ties, we established a common garden with individuals distance of pairs of populations was derived following Wright from three mesic forests in Hawaii that represent gradi- (1978). A genetic distance greater than zero indicates some degree of genetic differentiation among populations, and a distance of one ents in soil N and P availability. Large differences in nu- indicates complete divergence. trient availability among these sites could have created distinct selection pressures on the plants, particularly for traits related to nutrient use. We tested the populations Common garden setup occupying the different sites for intrinsic differences in In July 1996, 10–40 cm tall seedlings of M. polymorpha were col- tissue quality and growth rate. Finally, to assess the in- lected from fallen tree trunks, transplanted to 2-gallon pots, and fluence of genetic variation on any phenotypic variation placed in an open area at the University of Hawaii Volcano Exper- observed in field-grown plants, we compared the com- imental Farm in the Olaa Forest Reserve, Hawaii. A mixture of sand, vermiculite, and cinder was used as potting soil. Pots were mon garden results with in situ data obtained in earlier rearranged at random every 3 months to reduce effects of environ- experiments. mental heterogeneity within the garden site. Collection from nurse logs ensured that seedlings experienced a relatively common root- ing substrate across sites before transplant. Maternal effects on Materials and methods seedling phenotypes are likely to be a minimal source of variation, as seed sizes of M. polymorpha are small (about 0.1 mg; Corn Sites 1972). Response to nutrient availability was examined through N and Our study areas are located near Thurston lava tube in Hawaii Vol- P fertilization treatments in the common garden. Seedlings re- canoes National Park (Island of Hawaii), in Laupahoehoe Forest ceived either high or low levels of N and P in a factorial design. Reserve (Island of Hawaii), and in the Napali-Kona Forest Re- Plants in high N and P treatments received 6 mg P month–1 (as tri- serve (Kauai) (Fig. 1). These sites have soil substrates of different ple superphosphate) and 22.5 mg N month–1 (as ammonium ni- ages (300-, 20,000-, and 4.1 million years old, respectively), but trate), respectively. Low levels of N and P were 10% of high fer- are relatively uniform in parent material (volcanic tephra), eleva- tilization levels. Overall, at least ten individuals from the three tion (~1,200 m), precipitation (2,500 mm/year), average tempera- sites were assigned each of four possible treatments: high N/high ture (16°C), plant species composition, and topography [described P, high N/low P, low N/high P, or low N/low P. All plants received in detail by Crews et al. (1995) and Kitayama and Mueller-Dom- bois (1995)]. Phosphorus availability (as resin P) varies six-fold among the sites and is highest at Laupahoehoe, while N availabili- ty (as resin NH4–N and NO3–N) varies four-fold and is highest in Kauai (Table 1, Crews et al. 1995). Long-term fertilization experi- ments at these sites demonstrate that plant productivity is limited by N at Thurston (Vitousek et al. 1993), P at Kauai (Herbert and Fownes 1995), and by neither N nor P alone in Laupahoehoe (Vitousek and Farrington 1997). We will therefore refer to Thurs- ton, Laupahoehoe, and Kauai as N-limited, fertile, and P-limited sites, respectively. Genetic analyses To determine whether the populations of M. polymorpha had dif- ferentiated from one another genetically, we analyzed allozymes on individuals from each population and calculated genetic dis- tance among populations. Eight to ten individuals from each popu- lation were scored for genotypes at ten loci encoding eight func- Fig. 1 Locations of chronosequence sites across the Hawaiian Is- tional groups of enzymes (Table 2). We followed protocols devel- land chain Table 1 In situ phosphorus and nitrogen availability. Resin N is the sum of resin
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