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Rozzi Et Al 1997 Plant Ecology.Pdf Plant Ecology 132: 171±179, 1997. 171 c 1997 Kluwer Academic Publishers. Printed in Belgium. Ecological factors affecting gene ¯ow between populations of Anarthrophyllum cumingii (Papilionaceae) growing on equatorial- and polar-facing slopes in the Andes of Central Chile Ricardo Rozzi1;2, Mary Kalin Arroyo1 &JuanJ.Armesto1 1Laboratorio de Sistematica y EcologÂia Vegetal, Facultad de Ciencias, Universidad de Chile, Casilla 653, Santiago, Chile; 2Present address: Department of Ecology and Evolutionary Biology, University of Connecticut, 75 North Eagleville Road, Storrs, Connecticut 06269±3042, USA Received 19 March 1996; accepted in revised form 28 April 1997 Key words: Anarthrophyllum cumingii, Andean ¯ora, equatorial-facing slopes, ¯owering periods, gene ¯ow, polar-facing slopes, pollen ¯ow, pollination, seed dispersal Abstract In the Andes of Central Chile, ¯owering commences 1±2 months earlier on equatorial-(north-) facing than on polar- (south-) facing slopes, and pollinator assemblages also differ between these habitats. In order to understand the potential in¯uence of these differences on gene ¯ow, we monitored ¯owering periods and insect visitation rates to ¯owers of 60 individuals of Anarthrophyllum cumingii (Papilionaceae) on two equatorial- and two polar- 0 0 facing slopes in the Andes of central Chile (33 35 S;70 32 W). Flowering began about 30 days earlier on equatorial-facing slopes. Flowering periods of individuals on slopes with the same aspect had a mean overlap of 0.52, while those on opposite slopes had a mean overlap of 0.15. On equatorial-facing slopes Yramea lathionoides (Lepidoptera) accounted for 60% of the visits to ¯owers of A. cumingii, while on polar-facing slopes Centris cineraria (Hymenoptera) was responsible for more than 80% of ¯ower visits. Average similarities of visitor assemblages among individual plants on slopes with the same aspect was 0.83, while the mean similarity between individuals on opposite slopes was only 0.23. Within slopes ¯uorescent dyes were dispersed up to 40 m from the donor plants, but there was no movements of dyes between individuals growing on opposite slopes, even when they were separated by less than 10 m. Synchronous blooming and a common pollen vector are necessary conditions for pollen exchange between individuals. The overall probability of pollen exchange estimated by multiplying the inter-individual overlap for both factors, was nearly 0.5 for individuals growing on slopes with the same aspect, and less than 0.04 for individuals growing on opposite slopes. Consequently, at equivalent distances, the probability of pollen exchange between individuals growing on slopes of opposite aspect is more than 10-times lower than between those growing on the same slopes. Seed dispersal cannot compensate for restricted gene ¯ow through pollination, because seeds of A. cumingii were dispersed less than 2 m away from a parent plant. Presumably, restricted gene ¯ow could enhance genetic divergence between populations on slopes of contrasting aspects. This factor could be important in contributing to the high diversity and endemism in the Chilean Andes. Introduction only occur between simultaneously ¯owering individu- als that share a pollinator (Handel 1983). Although Most studies on gene ¯ow on plant populations focus variation in ¯owering periods (e.g., Ashton 1975; on distance between individuals as the decisive factor Hodgkinson and Quinn 1978; Hoefs 1979; Primack (Levin & Kerster 1969, 1974; Schall 1975, 1980). 1980, 1985a, b; Herrera 1986; Fripp et al. 1986) and Other factors, such as differences in ¯owering peri- pollinators (e.g. Primack 1978; Linhart & Feinsinger ods or pollen vectors, could also affect gene ¯ow 1980; Price & Waser 1982) frequently occur between in plant populations because pollen exchange can populations or individuals of a single species, the in¯u- GR: 201002120, Pips nr. 139933 BIO2KAP *139933* veg10588.tex; 28/08/1997; 14:46; v.7; p.1 172 ence of these factors on gene ¯ow has rarely been ¯oristically rich association of low, often spiny shrubs, examined (Handel 1983). perennial herbs and abundant geophytes up to 3000 m In the Andes of Central Chile, populations of found above timberline, and known as the subandean many plant species that occur on equatorial- (north- scrub (Arroyo et al. 1983). The study was conducted on ) facing slopes begin their ¯owering earlier than their Anarthrophyllum cumingii (H. et A.) Phil. (Papilioni- conspeci®cs growing on polar- (south-) facing slopes aceae), a small shrub about 0.5 m2 in area and less than (Arroyo et al. 1981; Reader 1984; Rozzi et al. 1989). 0.2 m in height (Rozzi 1990), endemic to the Andes of Equatorial- and polar-facing slopes in Central Chile Central Chile (Soraru 1974). This is one of the dom- differ sharply in microclimatic conditions (Armesto & inant species occurring at similar densities on both MartÂinez 1978; Armesto & Gutierrez 1978; Armesto equatorial- and polar-facing slopes between 2200 m et al. 1979; del Pozo 1985; Rozzi et al. 1989). Because and 2900 m. It has orange-yellow solitary ¯owers, is insect distribution and activity are greatly in¯uenced partially self-incompatible (Rozzi 1990), and is pol- by temperature (Heinrich 1974, 1975, 1979; Fogle- linated only by insects (Arroyo et al. 1982). Legumes man 1982; Kingsolver 1983, 1985a, b; Pivnick & Mc are 2 cm long, containing 1 to 5 seeds (Soraru 1974), Neill 1986, Corbet 1990, Herrera 1995), these plant probably dispersed by gravity (van der Pijl 1972) over populations could also differ in their insect pollinators. short distances. Likewise, along altitudinal gradients in central Chile ¯owers in conspeci®c populations at different eleva- tions are visited by different insect species, and this Methods variation has been related to the decrease in temperat- ure with increasing altitude (Arroyo et al. 1982, 1985, Flowering phenology 1987, 1990). We set out to quantify differences in ¯owering Flowering periods for 15 individuals of Anarthrophyl- periods and pollinator species between populations of lum cumingii were recorded on each of the four slopes. Anarthrophyllum cumingii, a common shrub growing Every 3 or 4 days, we counted on all the open ¯owers on both equatorial- and polar-facing slopes in the high within two ®xed 10 10 cm areas on the shrub crown. Andes of Central Chile. We also estimated pollen and seed dispersal distances. Finally, we discuss the signi- Pollinators ®cance of ¯owering and pollinator differences for gene ¯ow between plant populations growing on opposite Flower visitors and their visitation rates to ¯owers were slopes. determinated by two observers simultaneously mon- itoring Anarthrophyllum cumingii individuals bloom- ing on each slope. Observations were usually made Study area between 9 am and 6 pm, over three alternate 10 min intervals every hour. For each time interval, we recor- We chose two adjacent pairs of slopes of opposite ded the number of ¯owers visited on each individual of aspect, located at 2500 m elevation in the Andes of A. cumingii and the observed pollinator species. In all, 0 0 Central Chile (33 35 S; 70 32 W). One pair consisted we observed 1872 ¯owers on 73 shrubs for 1300 min of slopes with northeast- or equatorial- (300 )and distributed among 43 h. To determine whether insect south- or polar- (180 ) aspects, with slope angle of visitors carried pollen of A. cumingii, we captured 25 27 and 30 , respectively. These slopes were separ- individuals of each species that visited more than 10% ated by a gorge approximately 100 m deep. The other of the ¯owers, and examined them under optical and pair of slopes had aspects of 360 and 180 , and slope scanning electron microscopes. angle of 22 and 29 , respectively, and were separated by a smooth depression, less than 3 m wide and 2 m Pollen ¯ow deep. Hence, some of the individual plants growing on opposite slopes were separated by less than 10 m. To estimate pollen dispersal distances, in 1987 and The area has a mediterranean- type climate (sensu 1988 we studied dispersal of ¯uorescent dyes (see di Castri & Hajek 1976) with some degree of contin- Waser & Price 1982). Each year, ¯uorescent dyes were ental in¯uence (Rozzi et al. 1989), and is covered with dusted on anthers of 50 ¯owers on two shrubs on each snow from June to September. Vegetation consists of a slope, using a different color for each shrub. Dyes were veg10588.tex; 28/08/1997; 14:46; v.7; p.2 173 placed on a total of 800 ¯owers of 16 A. cumingii indi- In February±March 1986 and January±February viduals. Over the two years, we collected a total of 1987, two groups of 30 Anarthrophyllum cumingii 1300 ¯owers from 52 shrubs at various distances from seeds were placed next to each of six conspeci®c the source plant and examined their stigmas under UV shrubs, to assess biotic dispersal. One group was placed light using a magnifying glass. under a wire cage with a 7 7 mm mesh allowing ants to pass but excluding birds and rodents. The other group Estimation of similarities was placed directly on the ground. We placed 50 g of crushed oats and 50 g of `canary-seeds' 25 cm away To calculate similarities in ¯owering periods and vis- from each group of A. cumingii seeds, to assess the itor fauna between individual plants, we used the Pro- presence of potential dispersal agents. portional Similarity Index: P = : jP P j ij ik s 1 0 5 Results (Colwell & Futuyma 1971), Flowering periods P = N =Y ij j ij . Flowering periods of the four Anarthrophyllum N For ¯owering periods, ij is the number of open cumingii populations extended from October to Janu- j i Y ¯owers on individual on date ,and j is the total ary (Figure 1). Flowering began in early October on number of open ¯owers counted among all census dates both equatorial-facing slopes, and 25 days later on j N on individual .
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