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Giants Invading the Tropics: the Oriental Vessel Fern, Angiopteris Evecta (Marattiaceae)

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Giants Invading the Tropics: the Oriental Vessel Fern, Angiopteris Evecta (Marattiaceae) Biol Invasions (2008) 10:1215–1228 DOI 10.1007/s10530-007-9197-7 ORIGINAL PAPER Giants invading the tropics: the oriental vessel fern, Angiopteris evecta (Marattiaceae) Maarten J. M. Christenhusz Æ Tuuli K. Toivonen Received: 26 October 2007 / Accepted: 26 November 2007 / Published online: 7 December 2007 Ó Springer Science+Business Media B.V. 2007 Abstract The Oriental vessel fern, Angiopteris species could be cultivated over a much wider range evecta (G.Forst.) Hoffm. (Marattiaceae), has its than where it currently is grown. The escape of native range in the South Pacific. This species has cultivated plants into nature is probably due to been introduced into other localities since the 18th distance from natural areas and is limited by local century and is now listed as an invasive species in ecological factors, such as soil conditions or compet- several regions (Jamaica, Hawaii and Costa Rica). itors. The predicted distribution in Asia and The purpose of our study is (1) to trace the Madagascar is similar to the native distribution of distributional history of the species, and (2) to model the entire genus Angiopteris. It can therefore be its potential future range based on climatic condi- assumed that most Angiopteris species have similar tions. The native range and the history of introduction climatic preferences, and the absence of A. evecta in are based on the existing literature and on 158 this predicted region may be due to dispersal specimens from 15 herbaria, together with field limitation. In the Americas there is no native observations. As there are taxonomic problems Angiopteris, but our climatic model predicts a vast surrounding A. evecta, we limited our analysis to potential habitat in tropical America; an invasion of samples from the Pacific, most closely resembling the A. evecta should be anticipated here in localities type from Tahiti. We modelled the potential range where the species is cultivated. Vessel ferns are using GARP species distribution modelling with known to alter the natural environment, which may basic climatic variables, elevation, and location in reduce local biodiversity. Since A. evecta is not yet relation to the coast. Analysis of past records shows widely cultivated, it is advisable to restrict the trade that the species is able to colonise new ecosystems and spread of the species and to discourage its with relative ease. The modelling reveals that the cultivation as an ornamental. The global climate data available for modelling is however not detailed enough to predict the spread of A. evecta on a local or regional scale. M. J. M. Christenhusz (&) Department of Biology, Section Biodiversity, University of Turku, 20014 Turku, Finland Keywords Alien flora Á Angiopteris evecta Á e-mail: maarten.christenhusz@utu.fi Caribbean Á Conservation Á Ferns Á GARP Á Hawaii Á Invasive plant species Á T. K. Toivonen Islands Á Jamaica Á Marattiaceae Á Pacific Á Department of Geography, University of Helsinki, P.O. Box 64, 00014 Helsinki, Finland Species distribution modelling Á Neotropics Á e-mail: tuuli.k.toivonen@helsinki.fi Vessel ferns 123 1216 M. J. M. Christenhusz, T. K. Toivonen Introduction Innis (2004) reported that the ability of this vine to rapidly overtop trees, fix nitrogen and emit isopro- Species introductions by humans to new areas have pene has a highly negative effect on natural nitrogen become more common due to increased transport, cycles, forest biodiversity, freshwater eutrophication trade, travel, tourism, and the demand for exotic new and local air quality. species for horticulture. All of these provide path- The giant hogweed (Heracleum mantegazzianum ways for organisms to cross biogeographical borders Sommier and Levier, Apiaceae), a species originally that they would never be able to cross on their own from the Caucasian mountains, has spread throughout (Vitousek et al. 1996). Some introduced species Europe as a garden plant. According to Pysek and become invasive and have negative effects on the Pysek (1995), this species can invade a variety of local biodiversity, economy and even public health vegetation types, although disturbed habitats are (Mack et al. 2000; Forseth and Innis 2004; Reaser more easily invaded. The same study also showed et al. 2007). Globally, biological invasion by alien that vegetation invaded by H. mantegazzianum has a species is a major threat to natural ecosystems and much lower species diversity than similar adjacent biodiversity. An alien plant becomes invasive if it is vegetation that has not been invaded; many herbs and able to maintain its population spread and take over shrubs are unable to survive when overshadowed by suitable habitats from native plants (Williamson the large leaves of this Heracleum. 1996). In the tropics various alien plants are found as Exotic plant species have been introduced both well, which invade natural ecosystems in a similar intentionally, as garden or crop plants, and uninten- fashion, altering local ecosystems and reducing tionally, with agricultural practices or the moving of biodiversity; an example is Miconia calvescens soil. The effects of an introduction on the ecosystem DC. (Melastomataceae), which forms dense stands are usually irreversible (Wittenberg and Cock 2001). crowding out the native flora of the Pacific There are various ways in which invasive plants can islands (Meyer and Florence 1996). Bamboo alter their environment; these are summarised by (Bambusa vulgaris Schrad., Poaceae), the rose apple Reaser et al. (2007). Approximately 39–64% of (Syzygium jambos (L.) Alston, Myrtaceae) and the invasive plant species alter the hydrology and African tulip tree (Spathodea campanulata Pal., biochemistry of the natural ecosystem (Gordon Bignoniaceae) are well known examples of species 1998). These may negatively affect native species, invading tropical forests. Their slowly decomposing resulting in a lower local biodiversity. In Finland, for leaf litter prevents the regeneration of native instance, vegetational formations invaded by lupine species, and may change nutrient cycles. The dense (Lupinus polyphyllus Lindl., Fabaceae) were found to thickets formed by bamboo and rose apple prevent have a substantially lower diversity of both plants and the penetration of native species (O’Connor et al. butterflies than adjacent vegetations (Valtonen et al. 2000; Bito 2007). 2006). Valtonen et al. (2006) also showed that Even though most invasive plants are seed plants, invasive plants could affect the diversity of higher a few alien ferns are also known to be invasive and to trophic levels as well, especially when invading present serious problems for conservation. Examples species occur in high densities. of invasive ferns are Angiopteris evecta and Cya- Some invasive species, such as the infamous thea cooperi (F.Muell.) Domin, both from the South Japanese knotweed (Fallopia japonica (Houtt.) Pacific, but now forming major infestations on the Ronse Decr., Polygonaceae), alter ecosystems by Hawaiian Islands, threatening the endemic flora forming dense stands, crowding out native species. (Wilson 1996). In this article we model the potential This knotweed was introduced into Europe as an distribution of one of these, known variously as ornamental in the 19th century (Christenhusz and ‘Oriental vessel fern’, ‘mule’s foot fern’, ‘giant fern’ Uffelen 2001), and soon became a pest. It has a broad or ‘king fern’: Angiopteris evecta, a species native to ecological tolerance and invades various habitats. An New Guinea, coastal Australia, Melanesia, Microne- even more dramatic invasion was the introduction of sia, and Polynesia (Rolleri 2003). This species was kudzu (Pueraria lobata (Willd.) Ohwi, Fabaceae) to brought into cultivation soon after it was discovered North America to prevent soil erosion. Forseth and (Forster 1786); it is now known to be invasive in 123 Giants invading the tropics 1217 several tropical regions, most notably in Hawaii, Material and methods Jamaica and Costa Rica (http://plants.usda.gov/ &nbsp, and http://www.hear.org). Data collection Ferns of the genus Angiopteris are of evolutionary interest, because they are eusporangiate and have a Distribution data for A. evecta were obtained from fossil history dating back to the Jurassic (Konijnen- several sources. We have identified 141 specimens of burg-van Cittert 1975). In addition, the plants are of A. evecta in 15 herbaria (BISH, CANB, CR, DUKE, H, ornamental value, due to their large rhizomes and HVNP, IJ, MO, MT, NSW, NY, P, TUR, UCWI, US; gigantic leaves. For these reasons they are commonly herbarium acronyms according to Holmgren and cultivated in (botanical) gardens worldwide. The Holmgren 1998), and this data set was supplemented genus consists of some 200 microspecies, occurring with 17 field observations. Specimen images and throughout (sub-) tropical Asia, the Pacific and information from the Global Biodiversity Information Madagascar (Mabberley 1997). Facility (http://www.gbif.org/) and the database of the Angiopteris evecta is a species with large, globular Hawaiian Ecosystems at Risk project (http:// rhizomes that can grow up to 1.5 m tall and about www.hear.org/) were evaluated, 35 of whose records 1 m across. The starchy rhizomes are sometimes were included in our database. A total of 158 records consumed or used to perfume coconut oil (Mabberley included 128 from Australia, the Cook Islands, Fiji, 1997), which may have led to early cultivation in French Polynesia, Guam, Indonesia (Papua), the Ma- tropical Asia. The enormous bipinnate leaves sprout riana Islands, Micronesia, New Caledonia, Niue, spirally from the rhizome and can grow up to 6 m Palau, Papua New Guinea, Samoa, the Solomon long and 3 m wide. At the base of every petiole there Islands, Tonga and Vanuatu, where the species is are fleshy stipules, which have proliferous buds that native, and 30 from Costa Rica, Hawaii, Indonesia can grow into new plants when the stipules break off (Java, Ambon) and Jamaica, where the species is and find suitable ground (Uffelen 1994; Hoshizaki introduced (Fig. 1). All records were georeferenced and Moran 2001). This vegetative reproduction using maps, Google Earth (http://www.earth.
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