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Kalanchoe × Houghtonii (Crassulaceae) As an Invasive Plant: Potential Distribution by Ecological Niche Modelling (ENM)

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Kalanchoe × Houghtonii (Crassulaceae) As an Invasive Plant: Potential Distribution by Ecological Niche Modelling (ENM) Kalanchoe × houghtonii (Crassulaceae) as an invasive plant: potential distribution by Ecological Niche Modelling (ENM) Botanical Institute of Barcelona MASTER’S DEGREE IN ECOLOGY, ENVIRONMENTAL MANAGEMENT AND RESTORATION Author: Pilar Cachón Supervisors: Jordi López-Pujol, Sergi Massó, Daniel Vitales Tutor (UB): Emilia Gutiérrez University of Barcelona October 2017 Kalanchoe × houghtonii (Crassulaceae) as an invasive plant: potential distribution by Ecological Niche Modelling (ENM) Botanical Institute of Barcelona MASTER’S DEGREE IN ECOLOGY, ENVIRONMENTAL MANAGEMENT AND RESTORATION Author: Pilar Cachón Supervisors: Jordi López-Pujol, Sergi Massó, Daniel Vitales Tutor (UB): Emilia Gutiérrez University of Barcelona October 2017 Contents ABSTRACT .................................................................................................................... 1. INTRODUCTION ................................................................................................... 1 1.1. Objectives ....................................................................................................... 3 2. MATERIAL AND METHODS.................................................................................. 3 2.1. Study species ................................................................................................. 3 2.2. Localities gathering and georeferencing ......................................................... 4 2.3. Environmental variables and human footprint ................................................. 5 2.4. Niche modelling .............................................................................................. 7 3. RESULTS .............................................................................................................. 9 3.1. Niche modelling and variable contribution ....................................................... 9 3.2. Potential distribution at present ..................................................................... 10 3.3. Potential distribution under future climate (2070) .......................................... 11 4. DISCUSSION ...................................................................................................... 12 4.1. Current distribution and potential distribution under the present climate ....... 12 4.2. Potential effects of future climate change on distribution .............................. 14 5. CONCLUSIONS .................................................................................................. 17 ACKNOWLEDGEMENTS ........................................................................................... 18 REFERENCES ........................................................................................................... 19 ABSTRACT Alien species are currently considered one of the main direct threats to global biodiversity in our planet. Some alien plant species can become invasive and damage ecosystems, leading to negative effects on the local and global economy and society. Here, we have studied Kalanchoe × houghtonii D. B. Ward (Crassulaceae), a hybrid species artificially created by the horticulturist A. D. Houghton with gardening purposes in the 1930s. It is a result of the crossing between K. daigremontiana Raym - Hamet and H. Perrier and K. delagoensis Eckl. and Zeyh., two endemic species from Madagascar. Soon, the hybrid taxon showed large colonizing capacity, escaping outside the cultivation spots and rapidly expanding its distribution area. Now, K. × houghtonii is currently found in all continents except Antarctica. Therefore, considering the well-known background of the species, as well as its strong invading abilities, this plant represents an attractive model to investigate the expansion of alien species. The aims of the present study are evaluating the potential worldwide habitat of K. × houghtonii at different time frames, from present to future, employing various scenarios of climate change. To reach these goals, we first carried out a documentary research, involving the finding of localities where the species is present, through online databases, citizen science web portals, as well as many published academic papers. With the obtained occurrences, and using the maximum entropy algorithm implemented in MaxEnt, we modelled the potential habitat of the species to the present, which was later projected to the future under different scenarios. Results derived from this study would allow us to better understand the invading behavior of species with high colonizing potential such as K. × houghtonii, and, at the same time, inferring possible range contractions or expansions of the species across its whole distribution area under various scenarios of climate change. 1. INTRODUCTION Introduction and subsequent invasion of non-native species is the second leading cause of global biodiversity loss (Pyšek et al., 2004), resulting in altered ecosystem functions, increased vectors of diseases, and reduced distribution and diversity of native populations, which also means economic and health damages (Vitousek et al., 1996). A large proportion of invasive alien species worldwide were intentionally introduced to the areas where they are currently invasive, and many were widely disseminated once introduced to provide some value to humans (Ewel et al., 1999). A large proportion, if not the largest, of invasive plant species are introduced for horticultural use (Reichard and White, 2001; Kowarik, 2005). Ornamental plants are most likely to invade at the urban/wildland interface, where human habitation borders on natural vegetation. The cultivation and tending of plants in gardens produces high propagule pressure, so that many seeds and other propagules can spread into the surrounding natural vegetation. This then increases the likelihood and rate of any particular species invading natural and semi-natural ecosystems (Sullivan et al., 2004; Foxcroft et al., 2008). An example of a plant introduced due to ornamental uses that have become a widespread invader is Kalanchoe × houghtonii D. B. Ward, an artificial hybrid obtained by experimental crosses by the eminent horticulturist A.D. Houghton in the 1930s in his California greenhouses (Guillot et al., 2014). This species comes from the crossing of two of the species of the genus most frequent in cultivation, Kalanchoe daigremontiana Raym - Hamet and H. Perrier and K. tubiflora (Harv.) Raym.Hamet (Guillot et al., 2014). The invasion capability K. × houghtonii is enhanced by vigorous clonal growth through pseudobulbils (Figure I) that arise from the margin of their leaves (hence the popular name ‘mother of millions’ or ‘mother of thousands’), and this feature has been acquired through their progenitors (Guerra-García et al. 2015). Kalanchoe × houghtonii was firstly observed in the wild in Australia as early as 1965 (Guillot et al., 2014), and the plant has been reported throughout America (including the Caribbean islands), southern Europe, Asia (India, China), and Oceania (apart from Australia, in some Polynesian islands and in New Zealand) (Guillot et al., 2014; Wang et al., 2016). In some of these areas it has become a strong invader, such as in Queensland, Australia (Queensland Government, 2016) and Venezuela (erroneously identified as K. daigremontiana; Herrera et al. 2012). In southern Europe, such as in the Mediterranean coast of Spain, this hybrid is even more invasive than its parental species, Page 1 of 30 and in some cities it has already become a common component of the urban landscape (Guillot et al. 2014). From a management point of view, it is extremely important to identify areas which are not yet invaded but where early warning detection and control programs are essential to implement (Fang and Wan, 2009). Recent studies have developed niche- based models to assess the suitability of a region for a given species and its potential to spread throughout (Jimenez-Valverde et al., 2011). Ecological niche model using occurrence data and habitat environmental variables play important roles in predicting the potential distribution for alien plants (Thalmann et al., 2015). One of the most used methods of niche modelling is the maximum entropy algorithm as implemented in the software MaxEnt. It assesses the probability distribution of the study species by estimating the probability distribution of maximum entropy (Phillips and Dudík, 2008). The modelling is based on the combination of occurrence data with climatic layers and this method has the advantage that is based on only presence registers (Philips et al. 2006). Similarly, present-day distributions of species can be combined with environmental variables to enable projected distributions of species under future climate scenarios (Berry et al., 2002). a b Figure I. Kalanchoe × houghtonii. (Authors: a) S. Massó b) J. López-Pujol Page 2 of 30 1.1. Objectives The aims of this study are the following: (i) to explore the geography pattern of the distribution of Kalanchoe × houghtonii globally, gathering all the occurrences, both published in standard publications (e.g., articles and books) and available from other sources (e.g., databases including those based on citizen science, grey literature, personal communications, and even personal blogs), (ii) predicting the potential distribution of the species based on the gathered occurrences; such potential worldwide distribution will be predicted using the maximum entropy algorithm implemented in the software MaxEnt; and (iii) predicting how the distribution of this hybrid taxa is going
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