Natural Interspecific Hybridization in Gunnera (Gunneraceae) of the Juan Fernandez Islands, Chile!

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Natural Interspecific Hybridization in Gunnera (Gunneraceae) of the Juan Fernandez Islands, Chile! Pacific Science (1991), vol. 45, no. 4: 389-399 © 1991 by University of Hawaii Press. All rights reserved Natural Interspecific Hybridization in Gunnera (Gunneraceae) of the Juan Fernandez Islands, Chile! PATRICIA PACHECO/,3 Too F. STUESSY,2 AND DANIEL J. CRAWFORD2 ABSTRACT: Natural interspecific hybridization between Gunnera bracteata and G. peltata (Gunneraceae) in the Juan Fernandez Islands, Chile, is analyzed morphologically and chemically. Parental types from isolated populations were compared with parents and intermediates occurring together in Quebrada Villagra on Masatierra. Two transects were made in that area, one in a relatively undisturbed site, and another along a disturbed path. Hybrid indexes and distance diagrams were used to analyze morphological relationships, and leafflavonoids revealed chemical affinities. Minor flavonoid divergence between parental species precluded detailed analysis of dynamics of hybridization. Morphological analysis revealed intermediacy in both transects, with intergrada­ tion back toward both parents. It is suggested that introgressive hybridization is occurring in Quebrada Villagra between these two wind-pollinated species, with more hybridization taking place in disturbed regions. Reduction in surface area and changes in the ecology ofMasatierra during the past four million years may have brought the two species into closer contact and aided hybridization. NATURAL HYBRIDIZATION IS A frequent phe­ tion can have an evolutionarily significant nomenon among flowering plants (Knobloch effect if it is followed by establishment of 1972, Grant 1981). Although hybrids and advanced hybrid generations, with or without hybrid swarms have been detected in nature in polyploid y, or by introgression (Grant 1981). man y plant groups (Tanowitz and Adams Although the breakdown of isolating mecha­ 1986, Do yle and Doyle 1988, Grant and nisms is often the first factor in the appearance Wilken 1988, Tortosa 1988), the role of of natural interspecific hybrids, the environ­ hybridization in plant evolut ion is still con­ ment determines whether these hybrids, ad­ troversial (Levin 1979). Hybridization on one vanced generation hybrids , or introgressants hand may be rejected as an evolutionarily may become established successfully (Stebbins important process because of low fitness and 1950). In a stable environment, hybrid s are at sterility ofthe hybrid derivatives. On the other a disad vantage with respect to parental popu­ hand , hybridization may be seen as an impor­ lations because all niches are filled and no tant process in the evolution of plant groups " hybrid" habitats exist where the hybrids can because it can be a source of recombination survive (Anderson 1948). Disturbance of the and therefore its importance can be as great environment disrupts stable habitats, creating as mutation and recombination (Grant 1981 ). new open areas and new ecological niches that A more realistic perspective is that hybridiza- can be exploited by hybrid derivatives. In these new niches, hybrid individuals, with an enriched gene pool (Crins et al. 1988), have 1 This paper represents a portion of a thesis submitted by the senior auth or in part ial fulfillment of the require­ higher fitness than the parental populations ments for the Ph.D. degree at Ohio State University. and can compete successfully with them. Dissertation support provided by NSF grant INT­ Oceanic islands, because of their recent 8317088. Manuscript accepted I August 1990. origin, offer new habitats within a restricted 2 Department of Plant Biology, Ohio State University, 1735 Neil Avenue, Columbus , Ohio 43210. geographical area. Hybridization might be 3 Present address: Departamento de Biologia, Univer­ favored in insular areas where segregates can sidad de Taka, Taka, Chile. exploit these new habitats and therefore con- 389 390 PACIFIC SCIENCE, Volume 45, October 1991 KILOMETERS r-r-l o 200 400 Masafuera Masatierra 80 0 W \\Santa Clara +33 05 FIGURE I. Map showing the location of the Juan Fernandez Archipelago in southern South America. tribute to the evolutionary success of the a 69% endemism at the specific level (Skotts­ group. Speciation on islands apparently has berg 1956). been accompanied by little genetic divergence The most conspicuous example of natural at enzyme loci (Helenurm and Ganders 1985, interspecific hybridization in the Juan Fer­ Lowrey and Crawford 1985, Witter and Carr nandez Archipelago occurs in Gunnera (Gun­ 1988), and congeneric species are usually neraceae). This genus is represented by three highly interfertile (Gillett and Lim 1970,Gan­ endemic species: G. bracteata Steud. ex. J. ders and Nagata 1984, Lowrey and Crawford Benn., G. masafuerae Skottsb., and G. peltata 1985, Carr and Kyhos 1981, 1986). Hence the Phil. All three species belong to subgenus potential for natural interspecific hybridiza­ Panke (Schindler, 1905), which is the largest tion in floras of oceanic islands is great. group in the genus, with ca. 45 species distrib­ One set of oceanic islands, the Juan uted in South America and Hawaii (Schindler Fernandez Archipelago (Figure 1), is a small 1905, St. John 1946, Biloni 1959, Mora-Osejo group ofthree islands (Masatierra, Masafuera, 1978, 1984, Gomez 1983). Gunnera peltata is and Santa Clara), located at 33° S latitude endemic to Masatierra, where it grows from and 580 km west of the coast of Chile. Al­ 350 to 500 m. Gunnera bracteata is endemic to though its geographical position is not as the same island, but is found at slightly higher dramatically isolated as that of the Hawaiian elevations, from 400 m to the highest ridges Archipelago, the angiosperm flora does show at 600 m. Gunnera masafuerae is found on the Hybridization of Gunnera-PACHECO, STUESSY, AND CRAWFORD 391 younger island (Masafuera), where it grows Ninety-four individual plants ofG.bracteata from 100 m to 1400 m along streams and and G. peltata and intermediates were ana­ on the walls of canyons. Skottsberg (1922) lyzed from eight populations (Figure 2). Seven commented on a highly variable population specimens ofG.peltata are from three isolated of Gunnera growing in Quebrada Villagra pure populations: (1) a population in Plazoleta on Masatierra. He hypothesized that inter­ del Yunque (Stuessy et al. 66I6A); (2) a mediacy was due to hybridization between G. population in Quebrada El Pangal (Stuessy et bracteata and G. peltata but did not investi­ al. 6314);and (3) a population in a ravine from gate the situation further. Cordon Chifladores down toward Puerto There are other reports of hybridization in Frances (Stuessy et al. 6671). Four specimens Gunnera. Beutzenberg and Hair (1962) men­ of G. bracteata are from four isolated popula­ tioned a possible hybrid swarm between two tions: (1) Valle Colonial (Stuessy and Sanders species growing in New Zealand. Palkovic 5001H); (2) Cerro Damajuana (Stuessy (1978) suggested hybridization between G. and Sanders 5104); (3) Corrales de Molina insignis and G. talamancana in a disturbed (Stuessy, Crawford and Wiens 11306); and area in Costa Rica, and Mora-Osejo (1984) (4) Cerro Agudo (Stuessy and Garcia 11661). mentioned hybrids between three sympatric Eighty-three morphologically variable indi­ species of Gunnera in Colombia (G. atropur­ viduals are from an intermediate population purea, G. brephogea, and G. pilosa). in Quebrada Villagra. Two transects were set During an expedition to the Juan Fernandez in that locality (Figure 2). Transect 1 (24 Islands in 1984 we critically reexamined the individuals; Stuessy et al. 6494) ran from the highly morphologically variable population bottom of the ravine up through the forest of Gunnera in Quebrada Villagra that was from 430 m to 575 m. This habitat has mentioned by Skottsberg (1922). This vari­ remained almost unchanged and represents able population is ofconsiderable importance a relatively undisturbed environment. Tran­ because if it is the result of hybridization, it sect 2 (59 individuals; Stuessy et al 6481) would be the most conspicuous example of followed the trail up from the ravine at 145 m interspecific hybridization in the entire flora to Selkirk's Lookout at 555 m and represents of the archipelago. The objectives of this the disturbed habitat. Sampling was non­ paper, therefore, are to (1) use morphological random because an attempt was made to and flavonoid data to clarify whether hybrid­ cover variation seen. The largest leaf from ization does occur between G. bracteata and each plant was chosen for morphological G. peltata; (2) understand the nature and analysis. Samples for flavonoid analysis were dynamics ofthis hybridization; and (3) evalu­ taken at the same time from the same leaf. ate its evolutionary importance. Four to six scales (modified leaves that cover the apical meristem) were chosen for detailed study . MATERIALS AND METHODS Morphological Characters Sampling Fifteen characters that best discriminated Plant material and data were collected dur­ the two parental species were analyzed both ing expeditions to the Juan Fernandez Archi­ in the field and in herbarium material (Table pelago organized by the Departments of Bot­ 1). Only vegetative characters were utilized any of The Ohio State University (OS) and because no plant was in flower and only a Universidad de Concepcion (CONC), Chile, small proportion of individuals had fruits. during January-February 1980, 1984, and Each datum represents the mean value of 1990. Vouchers of the parental species are three or four individual measurements. Bio­ on deposit at OS, with
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