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Biogeographical Kinetics on an Island Volcano (Capelinhos, Azores)

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Biogeographical Kinetics on an Island Volcano (Capelinhos, Azores) Insect Conservation and Diversity (2012) 5, 358–366 doi: 10.1111/j.1752-4598.2011.00169.x Biogeographical kinetics on an island volcano (Cape- linhos, Azores): fast colonisation rates and dominance of arthropod exotic species 1,2 1 SIMONE FATTORINI and PAULO A. V. BORGES 1Departamento de Cieˆncias Agra´ - rias – CITAA (Azorean Biodiversity Group), Universidade dos Ac¸ ores, Angra do Heroı´smo, Portugal and 2Water Ecology Team, Department of Biotechnology and Biosciences, University of Milano Bicocca, Milan, Italy Abstract. 1. The aim of this paper was to investigate the arthropod colonisation of a recently erupted volcano in the framework of a general model of colonisation kinetics. 2. We analysed the diversity of arthropod communities at three locations on Faial Island (Azores) using a well-defined disturbance gradient: (i) a site that is new land added by the eruption of Capelinhos Volcano of 1957; (ii) a site moderately affected by this eruption; and (iii) a pristine site not affected. We calculated the recolonisation times at the disturbed sites using species richness at the undisturbed site as an equilib- rium value (last erupted 900–1000 years ago). 3. Species with different distributional ranges (endemic, native non-endemic and introduced) have different colonisation kinetics. Introduced exotic species were par- ticularly rapid in colonising the erupted volcano, reaching a number of species greater than that observed in the undisturbed area. By contrast, native non-endemic species had more difficulty in recolonising the erupted area, and no endemic has reached it. The volcano community is dominated by a few species with high abun- dance and shows low richness and strong dominance in comparison with the undis- turbed community. The moderately disturbed site supports a rich and well-balanced arthropod community. 4. Although the erupted volcano has a species richness even slightly higher than the undisturbed site, this is a consequence of the high colonisation ability of intro- duced species, and its arthropod community is strongly disharmonic. Key words. Azorean Islands, community ecology, ecological succession, equilib- rium theory, recolonisation, volcanic eruption. Introduction librium value, corresponding to the saturation of available niches (see Whittaker & Ferna´ ndez-Palacios, 2007). Earlier evi- Each archipelago has its own colonisation and evolutionary his- dence of saturation in animal assemblages was reported for tories that filter species both arriving at long-distance dispersal island birds in the Caribbean (Terborgh & Faaborgh, 1980) and and brought accidentally by man (Ricklefs, 2008). Classical fish assemblages in lakes (Tonn et al., 1990), but many more models of island biogeography postulate that insular species examples can be listed (reviewed by Srivastava, 1999). More assemblages are random samples drawn from a pool of potential recently, Winkler and Kampichler (2000), Soares et al. (2001) colonists and that species richness increases with time to an equi- and Borges and Brown (2004) found saturation patterns at a local scale, respectively, in grassland Collembola, litter ant com- Correspondence: Simone Fattorini, Water Ecology Team, munities and pasture spiders. However, this could indicate Department of Biotechnology and Biosciences, University of Mi- pseudo-saturation (i.e. the appearance of saturation by an unsat- lano Bicocca, Piazza della Scienza 2, I-20126 Milan, Italy. urated community) owing to small-scale effects and uncertainty E-mail: [email protected] in local species richness estimates (Caley & Schluter, 1997), and Ó 2011 The Authors 358 Insect Conservation and Diversity Ó 2011 The Royal Entomological Society Island volcano recolonisation 359 recent studies on local-regional processes show that most local 2011), studies based on more robust data may be particularly communities are not saturated (see Loreau, 2000). For example, useful to elucidate (re)colonisation processes. thereisevidencethatisolatedoceanicislandsmaybenotsatu- In this paper, we take advantage of recent standardised sam- rated with species, especially the recent ones, if there was not suf- pling on the Azorean Islands (see, e.g., Borges et al., 2005, 2006) ficient time for a complete colonisation (Borges & Brown, 1999; to investigate the arthropod colonisation of a recently erupted Borges & Hortal, 2009), the empty niche space being filled in his- volcano. We will address some of the shortcomings of previous torical times by recently introduced species (see e.g., Schaefer works and add further momentum to the development of a gen- et al., 2011, showing that introduced plant species are more eral model of colonisation kinetics. Because of the lack of abun- likely to become invasive in the absence of closely related species dance data, all conclusions drawn by Fattorini were based only in the native flora of the Azores). on species richness. Here, we also used abundance data to test As a consequence of the dispersal difficulties imposed by the his predictions on the evolution of community structure during isolation of many remote volcanic archipelagos, as well as the colonisation. destructive influence of volcanic activity and, more recently, of human activities, most islands are probably in a non-equilibrium condition (see Whittaker, 1995). Therefore, regional processes Materials and methods are critical to local community assembly (Ricklefs, 2008), and the rate of saturation will vary according to the characteristics of Study sites the system under study (dispersal ability of the species, isolation, habitat diversity, etc.). We analysed the arthropod recolonisation of Capelinhos Vol- In a recent paper, Fattorini (2010) used the approach devel- cano, on the Azorean island of Faial. Faial is a young island oped by Diamond (1972) in the framework of the equilibrium (0.7Ma)ofsmallsize(172km2) dominated by a central large theory of island biogeography of MacArthur and Wilson (1967) caldera (1043 m above sea level – ‘Caldeira do Faial’) and to investigate the recolonisation kinetics of island and mainland located in the Azorean microplate. Several cinder cones are volcanoes. The main critical point in that paper was the estima- located on the sides of the Caldeira do Faial. Capelinhos Vol- tion of the equilibrial value of species richness, i.e. the expected cano is the most recent volcano from the Azores (1957–1958), value of species richness for the erupted volcano at the end of exploding dramatically in 1957 with a devastating underwater the recolonisation process, when ecosystems will be completely eruption. The volcano had two main phases, one explosive restored and equilibrium reached. Studies on volcanic eruptions (phreatomagmatic) and the second Strombolian, i.e., an activity are retrospective in nature, because data on plant and animal with low explosivity and affecting only the area closest to the communities on the volcanic area before the eruption are lack- volcano (see Forjaz, 2007 for details). As a result of these erup- ing, and in case of volcanoes emerged from the sea, it is obvious tions, a large amount of ashes of the first phase covered the wes- that there were no communities before eruption. Thus, to esti- tern part of Faial and a total of 2.5 km2 of new land was added mate characteristics of communities at equilibrium, one must to the island (Machado et al., 1962). refer to a pristine (undisturbed) site which is considered an equi- The Capelinhos Volcano is a relatively small volcano within librium counterpart of the erupted volcano (see Thornton, an island. From this respect, it differs considerably from the two 2007). For island volcanoes, i.e., volcanic islands that were main types of volcanoes discussed by Fattorini (2010): volcanoes almost completely devastated by recent eruption, Fattorini on mainlands, and island volcanoes, i.e., islands that are entirely (2010) used species richness from adjacent islands of similar area constituted by a large volcano and which are virtually sterilised not affected by recent eruptions, while for a mainland volcano, by volcanic eruptions. In the case of Capelinhos Volcano, the he calculated the expected value at equilibrium from sites of dif- volcanic eruption had some major impact in the western part of ferent size and locations using a species–area relationship. These the island of Faial, but other places remained more or less undis- estimates are subject to several possible source of errors: (i) turbed. This particular condition allowed us to eliminate some incompleteness of faunal inventories of the reference sites; (ii) possible confounding factors that affected previous research. problems attached to the use of parameters of the species–area The use of an undisturbed site on the same island of Faial as a function to account for differences in area size among sites; and reference for the equilibrium conditions avoided problems (iii) differences in the ecological settings of the areas used as equi- owing to differences in ecological settings which can affect com- librium reference because of their distance from the study site parisons between distant sites. Thus, in our ‘natural experiment,’ (e.g. differences in altitude, climate, etc.). These problems (which we can assume that all differences between Capelinhos Volcano are typical of most studies on volcano recolonisation) were dis- (treated site) and the undisturbed (control) site can unambigu- cussed in Fattorini’s paper, showing the limits to current data ously be attributed to the ‘treatment’ (i.e.,
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