Are island plant communities more invaded than their mainland counterparts?
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| Are island plant communities more invaded than their mainland counterparts?
Montserrat Vila, Joan Pino, Ana Montero & Xavier Font Abstract Questions: Are island vegetation communities more invaded than their mainland counterparts? Is this pattern consistent among community types? Location: The coastal provinces of Catalonia and the para-oceanic Balearic Islands, both in NE Spain. These islands were connected to the continent more than 5.35 million years ago and are now located o200 km from the coast. Methods: We compiled a database of almost 3000 phytosociological releves from the Balearic Islands and Catalonia and compared the level of invasion by alien plants in island versus mainland commu- nities. Twenty distinct plant community types were compared between island and mainland coun- terparts. Results: The percentage of plots with alien species, number, percentage and cover percentage of alien species per plot was greater in Catalonia than in the Balearic Islands in most communities. Overall, across communities, more alien species were found in the mainland (53) compared to the islands (only nine). Despite these differences, patterns of the level of invasion in communities were highly consistent between the islands and mainland. The most in- vaded communities were ruderal and riparian. Main conclusion: Our results indicate that para- oceanic island communities such as the Balearic Islands are less invaded than their mainland counterparts. This difference reflects a smaller re- gional alien species pool in the Balearic Islands than  Vila, M. (corresponding author: montse.vila@ebd. csic.es) & Montero, A. (anamontero@ebd.csic.es): Estacion Biologica de Donana (EBD-CSIC), Avda. Americo Vespucio s/n, Isla de la Cartuja, 41092 Sevilla, Spain.
Pino, J. (Joan.Pino@uab.es): CREAF (Center for Ecological Research and Forestry Applications) and Unit of Ecology, Department of Animal and Plant Biology and Ecology, Autonomous University of Bar- celona, 08193 Bellaterra, Spain. Font, X. (xfont@ub.edu): Plant Biology Department, University of Barcelona, Diagonal 645, 08028 Barcelo- na, Spain. in the adjacent mainland, probably due to differ- ences in landscape heterogeneity and propagule pressure. Keywords: alien plants; Balearic Islands; community similarity; Mediterranean communities; para-ocea- nic islands; releve; species richness. Nomenclature: Bolos & Vigo (1984–2001), Rivas- Martinez et al. (2001). Introduction Islands have been a focus of attention in the bio- logical invasion literature due to their high regional pool of alien species and the impact that invaders pose to native species and ecosystem processes (Elton 1958; Lonsdale 1999). However, several studies have challenged the hypotheses that islands are more sus- ceptible to invaders than continents (D’Antonio & Dudley 1994; Jeschke 2008), especially when com- paring their establishment and success of spread. Possibly, invasibility (i.e. intrinsic resistance of com- munities to invasion) does not differ between island and mainland communities but differences are due to differences in the history of invasion (Sol 2000). Recently a comparison of the level of habitat invasion across regions of Europe, including Great Britain, found a high inter-regional consistency in habitat invasion patterns. These results suggest that habitats are good predictors for invasion risk as- sessment independent of climate, human history and insularity status (Chytry et al. 2008a, b). While stu- dies have shown higher regional proportions of alien species pools (Lonsdale 1999) or higher occurrences of certain species (Gimeno et al. 2006) in islands compared to their mainland counterparts, there has been no rigorous analysis quantifying the differ- ences in alien species richness in such plant communities to date. Such a comparison should be conducted in paired homologous communities. Communities that are homologous to the mainland are often difficult to find in remote oceanic islands whose evolutionary histories and species composi- tions are very different from their mainland counterparts (Brown & Lomolino 1998). 
Mediterranean Basin islands, which are con- sidered to be hotspots for biodiversity conservation (Cowling et al. 1996; Medail & Quezel 1999), are at significant risk from invasive species, which are threatening endemic species, especially in coastal areas (Di Castri et al. 1990; Hulme 2004). Most Mediterranean islands are para-oceanic (i.e. they have been attached to the continent in the past) and share many floristic components with the adjacent mainland (Greuter 1995). They thus do not re- present a disproportionately low native species richness considering area–species relationships (Medail 2008). Therefore, these islands offer the opportunity to test whether island communities are more invaded than their mainland counterparts. We took advantage of the large datasets available on vegetation plots and compared the level of invasion in different community types from the Balearic Is- lands and the adjacent mainland in Catalonia (both in NE Spain). We aimed to answer the following questions: Are island vegetation communities more invaded than their mainland counterparts? Is this pattern consistent among community types? We hy- pothesized that island communities would be more invaded than their mainland counterparts, but pat- terns of invasion would be consistent between islands and mainland communities. Methods Study area The Balearic archipelago (5014 km2) is located about 200 km east of Spain (Fig. 1). These islands are considered to be para-oceanic as they were con- nected to the continent during the Messinian period (between 5.70 and 5.35 million years ago). The Balearic archipelago consists of four main islands (Mallorca, Menorca, Eivissa and Formentera) and a set of uninhabited small islands and inlets, in- cluding Cabrera and Dragonera, among others. Their geographical location is between 11110 and 41190 E and 401050 and 381380 N. The climate is typi- cally Mediterranean, with warm summers, mild winters and low precipitation. Climate variability is highly affected by island size and elevation range, which is highest in Mallorca and lowest in Formentera and Cabrera. Although the Balearic Islands encompass similar land uses and have predominantly calcareous soils, the relative propor- tions of these differ, as does the topography (Rita & Payeras 2006). Catalonia is located in the northeast of Spain, bounded in the north by the Pyrenees and on the east by the Mediterranean Sea. It is situated between 01150 E and 31150 E and 401300 N and 421400 N. Due
 Catalonia
   Balearic Islands
 25 0 125 km
Fig. 1. Study area in the Balearic Islands (883 plots) and in Catalonia (2053 plots). The shaded area in Catalonia (the inland province of Lleida) was not included for analysis. to its contrasting topography, geology and climate, the vegetation is very heterogeneous (Bolos & Vigo 1984–2001). For this analysis, only provinces with typical Mediterranean maritime climate and located at o120 km from the coast were considered. The Catalonia study area covered 19 946 km2 (Fig. 1). The landscape structure of Catalonia and the Balearic islands reflects the typical interaction be- tween man and climate in Western Europe and the Mediterranean region. Forest and shrubland dom- inate the hilly areas, as a result of agricultural land abandonment in the mid-20th century. In contrast, lowlands are intensively cultivated or urbanised. Most of the population and summer tourism con- centrates in coastal areas. Database
15 o5%; 1 5 5–10%; 2 5 10–25%; 3 5 25–50%; 4 5 50–75%; 5 5 475%. From the total BDBC releves, we selected those alliances having phytosociological classification and that occurred both in Catalonia (2053 releves) and in the Balearic Islands (883 releves). In total, 20 dif- ferent paired community types of Mediterranean distribution were selected (App. 1). Thus, extra- Mediterranean communities (i.e. Boreal, Atlantic, Continental) occurring in Catalonia but lacking in the Balearic Islands were discarded. From each releve we calculated the number of alien and native species (i.e. species richness) and the percentage of alien species. We also calculated the percentage cover of alien species, by assigning to each species the central value of the cover percen- tage range corresponding to its abundance– dominance code. A species was considered an alien if it originated in another region outside mainland Spain or the Balearic Islands and it was introduced accidentally or deliberately by humans after 1500 AD. Therefore, only neophytes were considered and archeophytes (i.e. alien species introduced before the 15th century) were excluded from analysis. The publication date of releves (plots, hereafter) considered in this study ranged from 1960 to 2003 (App. 1). It was not significantly different between islands and the mainland (F1, 2888 5 2.28, P 5 0.13) but it was significantly different between commu- nities (F19, 2888 5 11.31, Po0.0001). The interaction between island and mainland status and community was not significant (F1, 2888 5 1.14, P 5 0.30). Therefore, assuming the publication year as a sur- rogate for sampling period, this lack of interaction indicates that, on average, a particular community type was sampled during the same period in Catalo- nia and in the Balearic Islands. In fact, traditionally, many communities were sampled in parallel by the same school of botanists (Vigo 1998). Data analysis To overcome differences in sample size (i.e. number of plots), mean species accumulation curves were computed for alien species in the islands and on the mainland (Gotelli & Colwell 2001). For all communities pooled together and for each commu- nity type, the first-order Jackknife richness estimator was calculated based on 100 randomiza- tion runs. Mean accumulation curves and Jackknife estimates were computed using EstimateS vs. 8.0 (R.K. Colwell, http://viceroy.eeb.uconn.edu/ estimates). Differences in the occurrence of alien species (i.e. percentage of plots with any alien) between is- lands and mainland across communities were tested with a paired t-test. Differences in alien species richness, alien species percentage, alien cover per- centage and native species richness between islands and the mainland, and among communities were tested by a general linear approach with Type 3 sums of squares performed with STATISTICA (version 6, StatSoft Inc., Tulsa, OK, USA) after log transformation of native species richness data and square-root (0.51x) transformation for the other variables. Pair-wise differences between commu- nities were tested with a Tukey–Kramer test. Results Overall, 55 alien species were found in the da- taset (53 in Catalonia and only nine in the Balearic Islands). Two alien species in the Balearic Islands (Cynara cardunculus and Hedysarum coronarium) were not found in Catalonian plots. Total alien spe- cies richness was lower in the islands than on the mainland (Jackknife estimates in islands 5 10 ± 1 SD; on mainland 5 42 ± 2 SD; Fig. 2). Alien species richness per plot was also lower in the islands than on the mainland (Table 1). The overall occurrence of alien species was low. It was significantly higher in Catalonia than in the Balearic Islands (Table 1). Ruderal salty scrub and calcareous pseudo-steppe communities were never invaded. Besides these two communities, 10 of the other communities were never invaded in the Ba- learic Islands (Fig. 3). Of the seven alien species occurring in both the Balearic Islands and Catalonia, there were no sig- nificant differences between mainland and islands in percentage occurrence across plots (paired t-test 5 0.805, P 5 0.452). However, except for Oxalis pes- caprae and Agave americana, these species were  50 Catalonia
 45
40 35 30 25
15 10
0 1 3 5 7 9 11 13 15 17 19 21 Number of plots (x 102)
found in more community types in Catalonia than in the Balearic Islands (Table 2). Number of sampling plots was very unbalanced. However, there was no significant relationship be- tween the number of plots per community type and the mean percentage of alien species (Catalonia: Spearman rank correlation, r 5 — 0.75, P 5 0.45; Balearic Islands: r 5 — 1.28, P 5 0.20). The percen- tage of alien species was significantly higher in the mainland than in the islands (Table 1). The percen- tage of alien species was also significantly different between communities (X2 5 57.064, Po0.0001, Fig. 3). Ruderal annual forb communities were the most invaded, followed by tall-forb humid communities, ruderal tall grasslands and riparian galleries and thickets. No significant differences were found be- tween the other communities (Tukey–Kramer tests not shown). There was a significant interaction be- tween community type and mainland/island status (X2 5 184.224, Po0.0001), indicating that although most communities were more invaded in Catalonia than in the Balearic Islands, a few communities could be as invaded in the islands as in the mainland (e.g. tall-forb humid communities, ruderal perennial forb communities and riparian communities; Fig. 3). Alien percentage cover was also significantly dif- ferent between islands and the mainland (Table 1), and differences among communities followed the same patterns as for percentage of alien species. There was a positive correlation between the number of alien species per plot and alien plant cover (Cata- lonia: Spearman rank correlation, r 5 0.36, Po0.0001; Balearic Islands: r 5 0.38, P 5 0.03). However, there was a large variation in this relation- ship. In fact, a single alien species could range from o5% to 90% cover. In total, 1678 native species were found in the da- taset (1442 in Catalonia and 827 in the Balearic Islands). The number of native species per plot ranged from one to 80 species, and on average was larger in Catalonia (18.09 ± 0.20, mean ± SE) than in the Balearic Islands (15.23 ± 0.32; X2 5 907.161, Po0.0001). The number of native species per plot was also significantly different between communities (X2 5 15.567, Po0.0001). There was a significant in- teraction between community type and mainland/ island status (X2 5 242.483, Po0.0001), indicating that although, in general, native species richness was Species Balearic  Islands
Catalonia Statistic larger in Catalonia than in the Balearic Islands, in some communities the number of native species may
Alien occurrence 6.21 ± 2.42 16.23 ± 4.57 2.85, P 5 0.010 Number of aliens 0.04 ± 0.01 0.19 ± 0.02 588.56, Po 0.0001 Percentage of aliens 0.36 ± 0.08 1.38 ± 0.12 655.30, Po 0.0001 not differ, and in two community types it was even lower in Catalonia than in the Balearic Islands, Percentage of alien cover
0.40 ± 0.11 1.92 ± 0.23 271.57, Po 0.0001 namely ruderal perennial forb communities and cal- careous pseudo-steppe communities (Fig. 3).  40
Balearic Islands 35 Catalonia 30  25
20 15 10 5  0
 25
Balearic Islands Catalonia 20  15
10 5  0
Fig. 3. Number of native species and percentage of alien species per plot (mean ± SE) for different communities in Cata- lonia and the Balearic Islands. Communities follow the BDBC nomenclature (http://biodiver.bio.ub.es/biocat): L009 Reedbeds, L015 Hydrophytic vegetation, L021 Perennial humid meadows, L026 Chamaephytic vegetation of sea cliffs, L032 Brackish rushbeds, L035 Fixed dune dwarf scrub, L062 Ruderal annual forb communities, L063 Tall-forb humid commu- nities, L065 Ruderal grasslands, L068 Ruderal salty scrub, L073 Ruderal tall grasslands, L074 Ruderal perennial forb communities, L085 Trampled ruderal grasslands, L089 Siliceous pseudo-steppe communities, L092 Calcareous pseudo- steppe communities, L095 Xeric perennial grasslands, L0123 Calcareous Mediterranean scrub, L0125 Riparian galleries and thickets, L0147 Mediterranean macchia, L0148 Mediterranean sclerophyllous forests.
Contrary to the general wisdom that islands are more invaded than the mainland, we found that most plant communities in the Balearic Islands were less invaded than their mainland counterparts in Catalonia. This lower level of invasion is manifested
Table 2. Percentage occurrence across invaded plots of the seven alien species present in both the Balearic Islands and Catalonia, with indication of the invaded communities (in parenthesis), and reproduction and dispersal modes (super- indexs). V 5 vegetative, S 5 sexual reproduction; W 5 wind, Z 5 animal, U 5 unspecific seed dispersal. See Fig. 3 for community nomenclature.
by a lower occurrence of alien species, lower com- munity alien species richness and lower alien plant cover. However, there was a high consistency in the identity of the most invaded communities, which in general had intermediate levels of native species richness. These communities are located in nutrient- rich and intermittently wet areas. Invasion in such communities is consistent with the theory of fluctu- ating resource availability (Davis et al. 2000), which proposes that pulses in resource inputs or in their use by organisms (e.g. because of episodic perturbations) trigger the establishment and spread of invaders. The frequency of individual alien species across plots was low and did not differ between the islands and the mainland, indicating that the niche occu- pancy of alien species is small in both regions (Lambdon et al. 2008). On average, alien species in- vaded more community types in the mainland than in the islands, except for A. americana and especially O. pes-caprae. Both species have the advantage that they reproduce vegetatively. Extensive field surveys have also found O. pes-caprae invades more com- munity types in the islands than in the mainland, possibly because of higher dispersal opportunities in the islands than on the mainland due to soil transport and machinery sharing between agri- cultural fields (Gimeno et al. 2006). O. pes-caprae have a fast growth rate and very effective annual vegetative reproduction through underground bulbs, with a high sprout and establishment rate in a broad variety of environmental conditions (Vila et al. 2006, 2008). The low level of invasion in island communities compared to mainland communities cannot be at- tributed to spatial sampling bias. Even if sample size was smaller in the islands than on the mainland, re- sults from the species accumulation curves matched the observed alien species richness (Fig. 2). Further- more, communities with many replicates did not host higher levels of invasion. Mediterranean scler- ophylous forest and calcareous scrub in Catalonia best exemplify this lack of association, since they were the most represented in the database, with more than 300 plots each; however their level of in- vasion ranked very low. The lower level of invasion in the islands and also lower native species richness compared to the mainland can be related to a smaller regional pool of species in the Balearic Islands compared to Catalo- nia. In the Balearic Islands, the total number of alien species has recently been estimated to be 304 species (Moragues & Rita 2005), while in Catalonia it is around 450 species or even more, considering that the most up-to-date regional survey is 16 years older than in the islands (Casasayas 1989). The regional pool of native species in mainland East Spain ad- jacent to Balearic Islands (3134, Bolos & Vigo 1984– 2001) is also two-times larger than in these islands (1569, Rita & Payeras 2006). The primary causes of the lower level of inva- sion in the Balearic Islands than in Catalonia can be due to a less heterogeneous landscape and lower propagule pressure. Catalonia is a larger and more heterogeneous region than the Balearic Islands, and it therefore provides a greater diversity of habitats. Both total and alien species richness in Catalonia are related to habitat diversity at the regional scale (Pausas et al. 2003; Pino et al. 2005, respectively). On the other hand, differences in alien species rich- ness at the local scale are suggestive of differences in propagule pressure (Vila & Pujadas 2001). In fact, according to the National Statistics Institute (http:// www.ine.es), population density and the density of road networks in 2003 (the latest database sampling year) were larger in Catalonia (277.99 hab km — 2 and 0.60 km — 1) than in the Balearic Islands (181.54 hab km — 2 and 0.43 km — 1). These two an- thropogenic features are surrogates for propagule pressure (Lockwood et al. 2005; Chytry et al. 2008b).
These invasion patterns mirror other island studies that have found alien success to be more closely related to environmental variation, differ- ences in disturbance regimes and invasion history than to community invasibility (Sol 2000; Gabriel et al. 2001; Teo et al. 2003; Li et al. 2006). However, the Balearic Islands, although less invaded than mainland, might not be less vulnerable to invasion, provided that land-use changes and more opportu- nities for species dispersal occur (Sax & Gaines 2003).
Acknowledgements. We thank V. Pillar and two anon- ymous referees for their helpful comments. This study was partially financed by the Ministerio de Ciencia e Innova- cion projects CGL2007-61165/BOS, CSD2008-00040 CGL-2009-07515 and CGL2006-13421-C04-01, and the Junta de Andalucıa Excellence Project RNM-4031.
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Vila, M., Bartomeus, I., Gimeno, I., Traveset, A. & Moragues, E. 2006. Demography of the invasive geophyte Oxalis pes-caprae across a Mediterranean island. Annals of Botany 97: 1055–1062. Vila, M., Siamantziouras, A-S.D., Brundu, G., Camarda, I., Lambdon, P., Medail, F., Moragues, E., Suehs, C., Traveset, A., Troumbis, A.Y. & Hulme, P.E. 2008. Widespread resistance of Mediterranean island ecosystems to the establishment of three alien species. Diversity and Distributions 14: 839–851. Received 4 May 2009; Accepted 29 November 2009. Co-ordinating Editor: Dr. Valerio Pillar. App. 1. Main descriptors (mean ± SD) of plots across community types in mainland Catalonia and the Balearic Islands. The syntaxonomical nomenclature follows Bolos & Vigo (1984–2001) and Rivas-Martınez et al. (2001).     Code Alliance Community Number of plots Occurrence* Total alien species Publication year
L009 Phragmition australis Mainland Islands Mainland Islands Mainland Islands Mainland Islands Reedbeds 49 10 2.04 0.00 1.97 ± 0.97 0.00 ± 0.00 1986.76 ± 11.25 1983.30 ± 7.07
L015 Isoetion Hydrophytic vegetation 20 20 20.00 0.00 1.06 ± 0.01 0.00 ± 0.00 1990.60 ± 5.53 1990.00 ± 6.81 L021 Molinio- Holoschoenion vulgaris L026 Crithmo- Limonion
L035 Crucianellion maritimae L062 Chenopodion muralis Perennial humid meadows Chamaephytic vegetation of sea cliffs Brackish rushbeds Fixed dune dwarf scrubs Ruderal annual forb communities 96 35 1.04 8.57 1.98 ± 0.98 1.00 ± 0.00 1990.57 ± 11.75 1989.69 ± 6.23 145 93 6.21 0.00 2.99 ± 0.99 0.00 ± 0.00 1990.37 ± 9.90 1990.02 ± 8.81 47 21 8.51 0.00 2.00 ± 0.00 0.00 ± 0.00 1987.64 ± 10.15 1987.48 ± 7.37 58 17 20.69 0.00 10.94 ± 2.18 0.00 ± 0.00 1993.88 ± 6.30 1992.71 ± 3.53 65 14 86.15 28.57 57.35 ± 0.00 3.00 ± 0.00 1984.83 ± 17.17 1985.07 ± 8.31 L063 Silybo-Urticion Tall-forb humid communities 76 38 26.31 36.84 23.89 ± 3.00 8.89 ± 1.86 1986.12 ± 14.26 1986.74 ± 8.67
L065 Hordeion leporini L068 Salsolo- Peganion L073 Bromo- Oryzopsion miliaceae L074 Echio- Galactition L085 Trifolio- Cynodontion L089 Helianthemion guttati Ruderal grasslands Ruderal salty scrub Ruderal tall grasslands
pseudo-steppe communities 95 28 18.95 0.00 16.94 ± 2.58 0.00 ± 0.00 1988.64 ± 15.11 1988.93 ± 5.48
74 20 21.62 10.00 14.90 ± 3.48 2.95 ± 0.95 1997.04 ± 8.61 1990.90 ± 6.14 102 10 32.35 0.00 26.89 ± 7.16 0.00 ± 0.00 1988.50 ± 17.65 1988.80 ± 6.05
App. 1. (Continued).  Code Alliance Community Number of plots Occurrence* Total alien species Publication year
L092 Thero- Brachypodion L095 Brachypodion phoenicoidis L0123 Rosmarino- Ericion L0125 Rubo ulmifolii- Nerion oleandri L0147 Oleo- Ceratonion Calcareous pseudo-steppe communities Xeric perennial grasslands Calcareous Mediterranean scrub Riparian galleries and thickets Mediterranean macchia Mainland Islands Mainland Islands Mainland Islands Mainland Islands 202 71 0.00 0.00 0.00 ± 0.00 0.00 ± 0.00 1996.42 ± 7.54 1994.48 ± 3.44
66 52 6.06 3.85 4.96 ± 1.38 1.00 ± 0.00 1985.99 ± 13.50 1987.56 ± 6.64 363 135 0.83 0.00 1.00 ± 0.00 0.00 ± 0.00 1985.34 ± 14.60 1986.50 ± 8.14 23 10 21.74 20.00 8.82 ± 2.25 1.00 ± 0.00 1996.17 ± 7.86 1988.80 ± 6.61 121 222 0.83 0.90 1.99 ± 0.99 2.99 ± 0.99 1986.84 ± 16.64 1987.89 ± 8.84 L0148 Quercion ilicis Mediterranean sclerophyllous forest
*i.e. percentage of plots with any alien. 364 25 1.10 0.00 4.99 ± 1.40 0.00 ± 0.00 1991.04 ± 7.65 1991.52 ± 2.02 |