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Genetic Diversity and Differentiation of Invasive Acacia Longifolia in Portugal

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Genetic Diversity and Differentiation of Invasive Acacia Longifolia in Portugal Web Ecol., 18, 91–103, 2018 https://doi.org/10.5194/we-18-91-2018 © Author(s) 2018. This work is distributed under the Creative Commons Attribution 4.0 License. Genetic diversity and differentiation of invasive Acacia longifolia in Portugal Sara Vicente1,2, Cristina Máguas2, and Helena Trindade1 1Centro de Estudos do Ambiente e do Mar (CESAM), Centro de Biotecnologia Vegetal (CBV), Faculdade de Ciências da Universidade de Lisboa, Lisbon, 1749-016, Portugal 2Center for Ecology, Evolution and Environmental Changes (cE3c), Faculdade de Ciências da Universidade de Lisboa, Lisbon, 1749-016, Portugal Correspondence: Helena Trindade ([email protected]) Received: 30 January 2018 – Revised: 27 April 2018 – Accepted: 4 May 2018 – Published: 24 May 2018 Abstract. Acacia longifolia is an aggressive invader worldwide. This species was brought to Portugal by the forestry services and is now found throughout the country with very strong ecological and social impacts. Al- though several ecological and physiological studies have been carried out, molecular studies in this species are sparse. Particularly, genetic variability evaluation in invasive ranges clearly deserves more attention. The aim of this study was to evaluate the genetic diversity and the genetic differentiation among populations of the alien in- vasive A. longifolia under different Mediterranean conditions. For that we studied three sandy dune populations along the Portuguese coast: Osso da Baleia (mesomediterranean), Pinheiro da Cruz and Vila Nova de Milfontes (termomediterranean). All sampled acacia plants were located underneath a pine forest, with the exception of the latter case study, where we also sampled in a diversified habitat conditions associated with the margins of agriculture fields. According to the historical records, it was also possible to compare acacia samples from original plantations and recently invaded areas. We have used two molecular markers – inter-simple sequence repeats (ISSR) and microsatellites (SSR). ISSR analysis showed that A. longifolia populations at all sampled locations have similar levels of diversity and a relatively low differentiation (8PT D 0:135). However, samples tended to cluster according to the regional (macro-scale) collection site. Microsatellites confirmed this low dif- ferentiation pattern. Our results indicate that differences in phenology and reproductive success observed during a previous study cannot be accounted for by genetic diversity and differentiation alone. Furthermore, our results taken together with historical information of A. longifolia suggest a single original introduction and a subsequent acclimation process. 1 Introduction stand its invasive mechanisms in order to prevent transfer to adjacent soils (e.g. Marchante et al., 2008; Richardson The genus Acacia comprises several species considered to be and Rejmánek, 2011; Ulm et al., 2017). It is known that aggressive invaders, and the Australian species are consid- the invasive capacity of A. longifolia is due to its ability ered important subjects of ecological invasion studies due to to form a high number of root nodules in different soils their introduction to several regions worldwide (e.g. Brazil, and the establishment of a symbiotic relationship with the South Africa and Portugal), rendering them serious environ- soil’s mutualists (Rodríguez-Echeverría et al., 2008), includ- mental threats on a global scale (Richardson and Rejmánek, ing the genera Bradyrhizobium and Rhizobium (Rodríguez- 2011). Acacia longifolia (Andrews) Willd., also known as Echeverría et al., 2011; Birnbaum et al., 2012), enabling “Sydney Golden Wattle”, is a nitrogen fixing legume origi- spreading to nutrient-poor soils, such as the dune systems nal from south-eastern Australia and Tasmania. This species (Ulm et al., 2017). Also, A. longifolia promotes the suc- is one of the most aggressive invaders worldwide and has cess of its own offspring (Rascher et al., 2011) and has the been the subject of many studies which seek to under- Published by Copernicus Publications on behalf of the European Ecological Federation (EEF). 92 S. Vicente et al.: Genetic diversity and differentiation of invasive Acacia longifolia in Portugal ability to control available sunlight to other plants, eventu- new environmental conditions, and such local responsiveness ally outcompeting them and occupying their space (Rascher associated with some degree of plasticity, might be one of the et al., 2011). Furthermore, this species is capable of quick reasons for the invasion success of this species worldwide. regeneration after fires because seed germination is actually Despite all the studies mentioned above, there is limited promoted by the fire itself (Marchante et al., 2003). This is information concerning A. longifolia on a molecular level, in surely one of the reasons for its high invasiveness in Por- particular the genetic variation and development of genetic tugal, a country with a Mediterranean climate and frequent structure after introduction. In recent years, the interest of fires during the summer. molecular studies as a tool to elucidate genetic diversity for A. longifolia was introduced to the Portuguese coastlines invasive species has increased. When a species is introduced by the forestry services at different periods between the late to a new location, the environmental conditions it faces may 19th and 20th century as a way to prevent the erosion of be very different from the ones at its native location, and the sand dunes (Marchante, 2011). Since its introduction, this introduced species has to overcome a number of challenges species has invaded many adjacent environments including and adapt to the new environment (Schierenbeck and Ain- Pinus pinaster plantations, which has a high economic im- ouche, 2006). Furthermore, genetic diversity has great impor- pact (Fernandes et al., 2015). A. longifolia’s invasive success tance in the invasive capacity of a species, as was elegantly in Portugal is such that, in spite of its fairly recent introduc- suggested by Lavergne and Molofsky (2007). However, de- tion some 130 years ago, it can be found throughout the coun- spite its active role, genetic diversity might not be strictly try where it is negatively impacting the diversity of native necessary for invasion. Baker (1965) proposed that a species species (Marchante et al., 2003) and ecosystem functioning might be able to respond to different environmental condi- (Rascher et al., 2012; Ulm et al., 2017) such as changes in tions through phenotypic plasticity and little genetic evolu- plant diversity (Holmes and Cowling, 1997; Lorenzo et al., tion, coining the term “general-purpose genotype”, which 2012; Lazzaro et al., 2014), soil composition (Marchante has since then been the subject of many studies. et al., 2008; Souza-Alonso et al., 2015) and water availability Molecular markers are an important and useful tool in (Lorenzo and Rodríguez-Echeverría, 2015). molecular biology and can be used in studies of invasive Although there is a large distribution of this species un- species as means to identify source populations, determine der different conditions, its reproductive success is depen- invasive routes and elucidate invasive mechanisms and suc- dent on the site (Fernandes et al., 2015), in particular the cess (Handley et al., 2011). Many molecular markers have flowering period, since it influences fruit and seed produc- been used in population and phylogenetic studies in aca- tion (Pyšek and Richardson, 2008). A study on A. longifolia cias, including inter-simple sequence repeats (ISSRs), simple under two different environmental conditions – higher pre- sequence repeats (SSRs or microsatellites), chloroplastidial cipitation with lower average temperature (north of Portu- DNA and internal and external transcribed spacers (ITS and gal), and lower precipitation with higher average tempera- ETS respectively) (Thompson et al., 2012, 2015; Omondi ture (south of Portugal) – indicated large differences in the et al., 2010, 2016; Harris et al., 2012; Miller et al., 2011; Le phenology and potential invasiveness impacts due to signif- Roux et al., 2011, 2014; Fredua-Agyeman et al., 2008; Mu- icant differences in flowering success and consequent seed lumba et al., 2012; Josiah et al., 2008; Ndlovu et al., 2013; production (Fernandes et al., 2015). Under typical Mediter- Murphy et al., 2010; Brown et al., 2010, 2012). Further- ranean climate and sparser forest canopy, acacias showed more, several SSRs markers have been developed for Aca- a shorter flowering period when compared to those under cia species, such as A. atkinsiana (Levy et al., 2014), A. bre- the more mesic Mediterranean conditions and denser for- vispica (Otero-Arnaiz, 2005). A. dealbata (Guillemaud et al., est canopy. This resulted in differences in reproductive suc- 2015), Acacia (Senegalia) mellifera (Ruiz-Guajardo et al., cess, with acacias under mesic conditions being more suc- 2007), A. nilotica (Wardill et al., 2004), A. mangium (Butcher cessful (Fernandes et al., 2015). These results suggest an in- et al., 2000), A. senegal (also known as Senegalia senegal) fluence of local environmental factors in the phenology of (Assoumane et al., 2009). However, to the best of our knowl- A. longifolia, particularly the temperature and forest man- edge, SSR primers have not yet been described for A. longi- agement, which may cause anticipation in the phenological folia. events (Fabio et al., 2010). Later, Morais and Freitas (2015) In this study, we used two different
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