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The Essential Role of Tree-Fern Trunks in the Regeneration of Weinmannia Tinctoria in Rain Forest on Reunion,´ Mascarene Archipelago

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The Essential Role of Tree-Fern Trunks in the Regeneration of Weinmannia Tinctoria in Rain Forest on Reunion,´ Mascarene Archipelago Journal of Tropical Ecology (2007) 23:487–492. Copyright © 2007 Cambridge University Press doi:10.1017/S026646740700418X Printed in the United Kingdom The essential role of tree-fern trunks in the regeneration of Weinmannia tinctoria in rain forest on Reunion,´ Mascarene Archipelago Geraldine´ Derroire∗†, Laurent Schmitt∗, Jean-Noel¨ Riviere` ∗, Jean-Michel Sarrailh† and Jacques Tassin‡1 ∗ CIRAD, UMR 53, 7 chemin de l’IRAT, 97410 Saint-Pierre, Reunion,´ France † ENGREF, 14 rue Girardet, CS 14216, 54042 Nancy Cedex, France ‡ CIRAD, UPR 37, Campus International de Baillarguet, TA 10/D, 34398 Montpellier Cedex 5, France (Accepted 30 March 2007) Abstract: The regeneration processes and the spatial distribution of the shade-intolerant Weinmannia tinctoria seedlings in the south of Reunion´ were investigated, with a special focus on the role of tree-ferns in seedling establishment. Mature individuals and seedling patches of Weinmannia tinctoria, and also tree-ferns, were mapped on two 1-ha rain-forest plots. Weinmannia tinctoria has hemi-epiphytic behaviour since seedlings were generally located on plant supports (99.6%). Regeneration was better on a support than on the ground, and germination tests showed that seeds were light-stimulated. Tall tree-ferns (Cyathea spp.) were suitable supports. Cyathea glauca and C. excelsa were more suitable supports than C. borbonica. Weinmannia tinctoria establishment complies with the facilitation model of succession and is dependent on tree-ferns. Resum´ e:´ La reg´ en´ eration´ et la distribution spatiale des semis de Weinmannia tinctoria,espece` intolerante´ a` l’ombre, ont et´ e´ etudi´ ees´ dans le sud de La Reunion.´ Nous nous sommes interess´ es´ au roleˆ des fougeres` arborescentes sur l’emergence´ de plantules. Les adultes matures et les taches de semis (ensembles de plusieurs plantules) de Weinmannia tinctoria,de memeˆ que les fougeres` arborescentes, ont et´ e´ cartographies´ sur deux parcelles de 1 ha en foretˆ humide. Weinmannia tinctoria a un comportement hemi-´ epiphyte´ car les semis se situaient principalement sur des supports (99,6%). La reg´ en´ eration´ se rev´ elait´ meilleure sur les supports qu’au sol, et les tests de germination ont montre´ que les semences germaient mieux alalumi` ere.` Les grandes fougeres` arborescentes (Cyathea spp.) constituaient des supports privilegi´ es.´ Cyathea glauca et Cyathea excelsa representaient´ des supports plus favorables que Cyathea borbonica. W. tinctoria apparaˆıt donc comme une espece` qui s’installe principalement en utilisant le modele` de facilitation et qui depend´ de la presence´ de fougeres` arborescentes. Key Words: colonization, ferns, germination, islands, montane forest, regeneration, seedlings, succession INTRODUCTION (Brandani et al. 1988, Christie & Armesto 2003, Liao et al. 2003, Riera 1985). However, very few quantitative The high level of small-scale heterogeneity in rain studies deal with regeneration using live plant supports, forests contributes to the local coexistence of species that and only recent ones involve the role of tree-fern trunks share the same ecological niches in the adult stage but in seedling establishment (Coomes et al. 2005, Mehltreter have different regeneration niches (Grubb 1977). The et al. 2005). vegetation itself may create local conditions impacting Weinmannia tinctoria Sm. (Cunoniaceae) is a common on the regeneration of different species (Ben´ıtez-Malvido long-lived and emergent tree of the montane rain forest 2006, Male & Roberts 2005). The role of fallen trees of Reunion´ (Scott 1997). The species frequently occurs as on regeneration and seedling development has been juvenile and mature trees in late-successional stands. It highlighted in both temperate and tropical forests can therefore be expected that W. tinctoria uses specific regeneration mechanisms that allow it to regenerate in late-successional stands of Reunion´ montane rain 1 Corresponding author. Email : [email protected] forests. Weinmannia racemosa was observed regenerating 488 GERALDINE´ DERROIRE ET AL. on tree-ferns in New Zealand (Beveridge 1973, Wardle plant species are endemic to the Mascarene Archipelago 1966), but the relative role of such living supports in the (Cadet 1980). The montane rain forest extends from recruitment of Weinmannia has never been investigated altitude 800 m to 1900 m on the windward side and in a quantitative way. In this study, we show the role 1100 m to 2000 m on the leeward side (Strasberg et al. of tree-fern supports in the regeneration of W. tinctoria 2005). It is characterized by the richness of epiphytes in montane rain forest. Our work is based on the and the abundance of tree-ferns. The dominant trees three following questions: (1) What are the regeneration and shrubs of the native montane rain forest are patterns and the spatial distribution of W. tinctoria Aphloia theiformis, Doratoxylon apetalum (Poir.) Radlk., seedlings in rain forests? (2) Do the three native tree-ferns Homalium paniculatum (Lam.) Benth., Nuxia verticillata (Cyathea borbonica Desv., C. excelsa Swartz and C. glauca Lam., Syzygium cymosum (Lam.) DC. and Weinmannia Bory) have a decisive role on the species regeneration? mauritiana D. Don (Tassin et al. 2004). The dynamics of (3) How do tree-ferns of different species or size (diameter native vegetation have been poorly documented (but see and height) classes determine W. tinctoria regeneration? Strasberg 1995). Weinmannia tinctoria is endemic to Reunion´ and Mauritius and is a common and even sometimes METHODS dominant species in montane rain forests of Reunion.´ This tree species can reach 15–20 m in height and 1 m Studied vegetation in diameter. Fruits are capsules containing 2–10 small ellipsoidal seeds, 0.7 mm long, with loose pubescence ◦ ◦ Reunion(21´ S,55 30 E,2512 km2)isavolcanicislandof (Scott 1997). About 9500 seeds are contained in a single the Mascarene Archipelago in the south-western Indian gram. Ocean (Figure 1). The species richness of the native flora is rather poor, consistent with the island’s small size, but endemism is high, as 55% of the 486 flowering Study sites Our fieldwork took place at two sites: the Riviere` des Remparts and Bon Accueil (Figure 1). On each site, data were collected on a 1-ha square study plot. The site of the Riviere` des Remparts is located on a recent basaltic flow from the Piton de la Fournaise volcano. The mean altitude of the study plot is 1630 m. Annual rainfall is 2100 mm (1974–2003) and mean annual temperature is 13.6 ◦C (1974–2003). The site of Bon Accueil is located on volcanic material from the Piton des Neiges volcano. The mean plot altitude is 1200 m. Annual rainfall is 2000 mm (1958–2004) and mean annual temperature is 16.8 ◦C (1958–2004). Soils are andisols on both sites (Raunet 1991). Data collection The spatial distribution and the regeneration of W. tinctoria were studied on both plots. Each plot was divided into 100 quadrats measuring 10 m × 10 m to make data collection easier and more precise. Plots and quadrats were delimited using topographical thread and a bearing compass. Within each quadrat Qi,allW. tinctoria and tree-fern specimens were mapped in XiYi coordinates, in order to deduce XY coordinates within the whole plot. The precision of X and Y was 0.1 m. Their height (H) and diameter at breast height (dbh) were recorded for individuals with dbh ≥ 8cm.Wedefinedaseedlingasa young plant grown from a seed and not exceeding 20 cm Figure 1. Location of the study sites. tall, and a seedling patch as a group of seedlings growing Tree-ferns and regeneration of Weinmannia tinctoria 489 on the same individual support or, if located on the soil, also used on germination results to test the effect of light. having the same XY coordinates. All seedling patches of On each site and for different types of support, a KW test W. tinctoria located on plant supports or soil were mapped. wasalsousedbetweenthenumberofW.tinctoriaseedlings Within each patch, we counted the number of seedlings by tree-fern and the type of support to test the suitability and recorded the microsite type (i.e. plant support or soil). of the support. This test was used for each tree-fern species When the support was vegetation, we also recorded the (Cyathea borbonica, C. excelsa and C. glauca). The site effect species of the plant support. The mapping of seedling was tested using the same KW test between the number of patcheswasdonequadratbyquadrat.Weusedmeasuring W. tinctoria seedlings by patch and the site (Bon Accueil tapes and a bearing compass. Data collection was carried and Riviere` des Remparts). For Bon Accueil, a KW test was out in May 2003 at Bon Accueil site, and between July carried out between the number of W. tinctoria seedlings 2002 and April 2003 at Riviere` des Remparts. on a tree-fern and the height classes of a tree-fern and In the laboratory, germination tests were conducted to between the number of seedlings and the diameter class of check that W. tinctoria produced light-stimulated seeds. a tree-fern. Height classes used were < 1m,1–2m,2–4m They were performed in transparent Petri dishes placed and 4–8 m. Diameter classes were < 8 cm, 8–17.5 cm, in a germination cabinet at 15 ◦C (average condition on 17.5–27.5 cm, 27.5–37.5 cm, 37.5–47.5 cm and 47.5– the distribution area of W. tinctoria). Two light treatments 57.5 cm (adapted from diameter classes routinely used in were tested: under constant light (under an illuminance French forestry). KW test was processed using SYSTAT. of 2500 lx) and in the dark. Seeds were placed on sterilized sifted river sand. The sand was saturated with water using a sprayer. Each treatment was tested on four replicates RESULTS of 25 seeds. The number of seeds that had germinated was regularly recorded until no more germination was Weinmannia tinctoria seedlings were rarely isolated.
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