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Allelopathic Effects of the Chilean Matorral Shrub Flourensia Thurifera

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Allelopathic Effects of the Chilean Matorral Shrub Flourensia Thurifera Revista Chilena de Historia Natural 60: 57-62, 1987 Allelopathic effects of the Chilean matorral shrub Flourensia thurifera Efectos alelopaticos del arbusto de matorral Flourensia thurifera EDUARDO R. FUENTES, GUILLERMO A. ESPINOZA and GLORIA GAJARDO Laboratorio de Ecologia, Pontificia Universidad Cat61ica de Chile, Casilla 114-D, Santiago, Chile ABSTRACT This paper reports 4 types of evidence of the allelopathic effect of the drought-deciduous shrub F1ourensia thuri[era (Compositae) on forbs and shrubs. Field evidences include: (1) observed differences in grass abundances at neigh- boring sites with and without (Temoved) F. thuri[era; (2) measured grass abundances at various distances and directions from isolated F. thurifera shrubs; and (3) the results of experiments in which F. thurifera was cut and the ensuing increment in the relative grass densities monitored. Laboratory evidences pertain to relative germination rates of rye and shrub seeds when irrigated with water in which shrub leaves had been soaked. All four evidences point in the same direction, namely, that F. thurifera produces water-soluble chemical substances capable of inhibiting the germination of other plant species. In addition, the paper supports previous results regarding the lack of demonstrable allelopathic effects in some evergreen species of shrubs. Finally, the ecological significance of these results is discussed. Key words: Allelopathy, Forb densities, forb supression, germination rates. RESUMEN Este articulo informa sobre 4 tipos de evidencias que sugieren que el arbusto deciduo de verano F1ourensia thuri[era (Compositae) tiene efectos alelopaticos sobre hierbas y arbustos. Las evidencias de campo incluyen: (1) Observaciones de abundancia de hierbas en sitios vecinos con y sin F. thurifera; (2) mediciones de abundancia de pastos a distintas distancias y en varias direcciones a partir del centro de arbustos aislados de F. thurifera, y (3) resultados de experi- mentos donde se extrajo las plantas de F. thurifera y se midi6 el incremento posterior de la cobertura de hierbas. Las evidencias de laboratorio se relacionan con las tasas relativas de germinaci6n de centeno y semillas de arbustos. Ambos grupos fueron regados con agua en la cual se remojaron previamente hojas de distintas plantas lefiosas. Las evidencias sugieren que F. thurifera produce sustancias capaces de inhibir la germinaci6n de algunas especies. Adicionalmente, este trabajo apoya resultados previos relatives a la carencia de efectos alelopaticos demostrables en algunas especies de arbustos siempreverdes. AI final se discute el significado ecologico de estos resultados. Palabras claves: Alelopatia, densidad de hierbas, supresi6n de hierbas, tasas de germinaci6n, plántulas. INTRODUCTION segregation due to the inhibition produced by metabolites of one plant on another. Allelopa thy, the chemical inhibition of But surprisingly, neither for the rich flora seed germination and plant growth, can of the Southern Cape area (South Africa), produce important community patterns. nor for central Chile, the other two areas Allelopathy has been reported to occur with wet winters and dry summers, there and produce such patterns in three out have been yet any claims of this type of of the five world areas with a mediterra- interference. The only published work nean-type of climate. In California (Muller (Montenegro et al. 1978) testing for 1966), the Mediterranean Basin (De1enil allelopathy in the Chilean matorral, reports 1951, Guyot 1951, Ballester & Visitez that aqueous leaf extracts of five common 1979) and in Australia (del Moral et al. species of shrub failed to produce measur- 1978) there have been reports of spatial able effects on the germination or growth (Received 12 May 1986. Accepted 2 March 1987). 58 FUENTES ET AL. of A vena sativa. These laboratory experi- second type of experiment the shrubs ments were confirmed by experiences in were cut at the end of June of 1983 the field in which shrub litter added to (winter), before the grasses started their various plots also failed in generating annual growing season (see Montenegro distinct effects on the herbaceous cover. et al. 1978) and final measurements were The aim of this contribution is to taken in November of the same year, document our field observations and after annual plant growth was completed. laboratory experiments suggesting that The first type of removal experiment the native shrub Flourensia thurifera was done on two 50 x 30 m plots, one (Compositae) is capable of producing of which was on an equator-facing slope allelopathic effects, not only on common and had an almost complete cover by field forbs, but even on other native shrubs. F. thurifera. There were also a few cacti F. thurifera is a drought-deciduous, wind- (Trichocereus chilensis) present. The second dispersed shrub that is infrequently found plot, on a northwest-facing slope was in the area where Montenegro et al. (1978) covered by a mixture of F. thurifera and worked (coastal ranges at 330 S.), but Colliguaya odorifera. For each plot we further north and on some of the drier had a neighbouring control site of the interior slopes further east, it dominates same size. the landscape (Etienne et al. 1982, and The other removal experiments were E.F. personal observations). In addition done on 50 x 10 m plots located on a our paper will confirm the previous results north-facing slope covered by F. thurifera by Montenegro et al. (1978) regarding and some sparse T. chilensis individuals. the absence of demonstrable allelopathic Here we made an additional type of control effects in some of the evergreen shrubs which consisted of eliminating only T. they used. chilensis cover from one plot. All these experiments and field obser- vations were carried out approximately METHODS 60 km north of Santiago at a site (32° 53' LS, 7QO 43' W) located at about 750 a) Field: Casual observations suggested m above sea level. a negative correlation between cover b) Laboratory. Our experimental design by F. thurifera and common naturalized followed the classical lines of our prede- forbs (Tricetobromus sp., Chaetanthera cessors (Me Pherson and Muller 1969) moechioides, Bromus spp., Vulpia der- and basically consisted of irrigating seeds toenis, Carthamus lanatus; see also Keeley of various species with solutions obtained & Johnson 1977). We quantified these from soaking leaves of several shrub species. observations by measuring relative abun- By following the time course of such a dance of these forbs in: (1) areas with series and by comparing the germination F. thurifera and in neighbouring areas rate with that of controls (distilled water), (30 m) where F. thurifera had been com- the evidence for allelopathy is obtained. pletely removed at least 10 years before; Fresh leaves ( 150 g) collected by stripping and (2) along radial transects out of iso- them from the stems, and distilled water lated F. thurifera shrubs. Relative grass (2 1) were used to prepare the various abundances were measured by the number solutions. The leaves were left in the water of contacts along linear transects or by for 36 hours before the solutions were throwing a 30 x 30 em metalic grid. This sieved and used on the seeds. To prepare grid was thrown 30 times on each experi- the four experimental solutions, we used mental plot as well as on each one of the leaves of Quillaja saponaria, Lithraea controls. caustica, Colliguaya odorifera and F. We also made experiments in which the thurifera. Of these, the first three species F. thurifera canopy was cut 5 em above were also used by Montenegro et al. ( 1978). ground level and grass relative abundances The experimental solutions were used to were monitored throughout the season. irrigate Petri dishes with 40 seeds each. We made two types of removal , experi- These seeds belonged to each of the ments. In the first F. thurifera was cut following four species: Q. saponaria, L. in the spring of 1982 and again in winter caustica, Acacia caven (Leguminosae) or 1983, but the measurements were done common rye. Seeds of L. caustica and A. only in the second year (1983). In the caven were scarified with a concentrated ALLELOPATHY IN THE CHILEAN MATORRAL 59 sulfuric acid solution before the experi- Results of the radial transects out of F. ment. Rye was used as it was the most thurifera shrubs (Fig. 1) show that forb similar seed to grasses available at the density is non-increasing (r = .20, P > .05) time, and because it allowed comparison and low under the canopy, but increasing with the previous results by Montenegro (r = .90, P < .01) and higher at greater et a/. (1978). All experiments were carried distances, particularly beyond the canopy out under laboratory conditions (180C projection. Moreover, the number of room temperature and 8- I 0 h fluorescent contacts at the first meter away from the light). trunk (Fig. 1b) was higher on the uphill than on the downhill side of shrubs (t-test, P < .01), suggesting that inhibition is not RESULTS a product of competition but of water- soluble compounds leaching from the a) Field. Relative forb densities with shrubs shrubs. and where shrubs had been removed at Summarizing, in closed stands of F. least 10 years before our experiment, are thurifera, there are clearly fewer forbs shown in Table 1a. It can be seen that than in open spaces, and where isolated forb density where F. thurifera had been F. thurifera individuals occur, they tend removed was several times higher than with to have fewer forbs under them, than the shrubs, notwithstanding the seasonal similar places somewhat more distant from increment in the shrub covered plot. the canopy projection. 6 10 Ill b u Ill -" v c0 4 - u ....0 8 a c 0 0 ... u Cll ..0 2 6 E 0 :::1 - z ....Cll 0 .0 4 E ~ z 2 w}JF /D 0 4 8 12 16 20 24 28 Distance (categories) Fig.
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