Aim Large-bodied vertebrates often have a dramatic role in ecosystem function through herbivory, trampling, seed dispersal and nutrient cycling. The iconic Galápagos tortoises (Chelonoidis nigra) are the largest extant terrestrial ectotherms, yet their ecology is poorly known. Large body size should confer a generalist diet, benign digestive processes and long-distance ranging ability, rendering giant tortoises adept seed dispersers. We sought to determine the extent of seed dispersal by Galápagos tortoises and their impact on seed germination for selected species, and to assess potential impacts of tortoise dispersal on the vegetation dynamics of the Galápagos.Location Galápagos, Ecuador. MethodsTo determine the number of seeds dispersed we identified and counted intact seeds from 120 fresh dung piles in both agricultural and national park land. To estimate the distance over which tortoises move seeds we used estimated digesta retention times from captive tortoises as a proxy for retention times of wild tortoises and tortoise movement data obtained from GPS telemetry. We conducted germination trials for five plant species to determine whether tortoise processing influenced germination success. ResultsIn our dung sample, we found intact seeds from > 45 plant species, of which 11 were from introduced species. Tortoises defecated, on average, 464 (SE 95) seeds and 2.8 (SE 0.2) species per dung pile. Seed numbers were dominated by introduced species, particularly in agricultural land. Tortoises frequently moved seeds over long distances; during mean digesta retention times (12 days) tortoises moved an average of 394 m (SE 34) and a maximum of 4355 m over the longest recorded retention time (28 days). We did not find evidence that tortoise ingestion or the presence of dung influenced seed germination success.Main conclusions Galápagos tortoises are prodigious seed dispersers, regularly moving large quantities of seeds over long distances. This may confer important advantages to tortoise-dispersed species, including transport of seeds away from the parent plants into sites favourable for germination. More extensive research is needed to quantify germination success, recruitment to adulthood and demography of plants under natural conditions, with and without tortoise dispersal, to determine the seed dispersal effectiveness of Galápagos tortoises.

BackgroundPlant invasions are causing habitat degradation in Galapagos. Problems are concentrated on the four inhabited islands. Plants introduced to rural areas in the humid highlands and urban areas on the arid coast act as foci for invasion of the surrounding Galapagos National Park.Methodology/Principal FindingsHere we present results of the most comprehensive inventory to date of alien vascular plants in the inhabited areas of Galapagos. The survey was conducted between 2002 and 2007, in 6031 properties (97% of the total) on Floreana, Isabela, San Cristobal and Santa Cruz Islands. In total 754 alien vascular plant taxa were recorded, representing 468 genera in 123 families. Dicotyledons represented 554 taxa, monocotyledons 183, there were 7 gymnosperms and 10 pteridophytes. Almost half (363) of the taxa were herbaceous. The most represented families were Fabaceae (sensu lato), Asteraceae and Poaceae. The three most recorded species in the humid rural areas were Psidium guajava, Passiflora edulis and Bryophyllum pinnatum, and in the dry urban areas, Aloe vera, Portulaca oleracea and Carica papaya. In total, 264 (35%) taxa were recorded as naturalized. The most common use for taxa was ornamental (52%).Conclusions/SignificanceThis extensive survey has increased the known alien vascular flora of Galapagos by 257 species, giving a ratio of alien to native taxa of 1.57∶1. It provides a crucial baseline for plant invasion management in the archipelago and contributes data for meta analyses of invasion processes worldwide. A repeat of the survey in the future would act as an effective early detection tool to help avoid further invasion of the Galapagos National Park.

The distribution of resources and food selection are fundamental to the ecology, life history, physiology, population dynamics, and conservation of animals. Introduced plants are changing foraging dynamics of herbivores in many ecosystems often with unknown consequences. Galapagos tortoises, like many herbivores, undertake migrations along elevation gradients driven by variability in vegetation productivity which take them into upland areas dominated by introduced plants. We sought to characterize diet composition of two species of Galapagos tortoises, focussing on how the role of introduced forage species changes over space and the implications for tortoise conservation. We quantified the distribution of tortoises with elevation using GPS telemetry. Along the elevation gradient, we quantified the abundance of introduced and native plant species, estimated diet composition by recording foods consumed by tortoises, and assessed tortoise physical condition from body weights and blood parameter values. Tortoises ranged between 0 and 429 m in elevation over which they consumed at least 64 plant species from 26 families, 44 percent of which were introduced species. Cover of introduced species and the proportion of introduced species in tortoise diets increased with elevation. Introduced species were positively selected for by tortoises at all elevations. Tortoise physical condition was either consistent or increased with elevation at the least biologically productive season on Galapagos. Santa Cruz tortoises are generalist herbivores that have adapted their feeding behavior to consume many introduced plant species that has likely made a positive contribution to tortoise nutrition. Some transformed habitats that contain an abundance of introduced forage species are compatible with tortoise conservation.

Introduced species present the greatest threat to the unique terrestrial biodiversity of the Galapagos Islands. We assess the current status of plant invasion in Galapagos, predict the likelihood of future naturalizations and invasions from the existing introduced flora, and suggest measures to help limit future invasions. There has been a 1.46 fold increase in plant biodiversity in Galapagos due to alien plant naturalizations, reflecting a similar trend on islands elsewhere. There are 870 alien plant species recorded in the archipelago. Of evaluated species, 34% species have naturalized. Within this group are the invasive species (16% of evaluated) and the transformers (3.3% of evaluated). We show that, as expected, naturalized species have been present in the archipelago longer than non-naturalized species. We also find that a higher human-mediated propagule pressure is associated with a greater human population and with properties that have been settled longer. This, combined with the relatively recent introduction of most species, leads us to the conclusion that Galapagos is at an early stage of plant invasion. We predict that more species from the existing alien flora will find an opportunity to naturalize and invade as propagule pressure increases alongside rapid human population growth associated with immigration to serve the booming tourism industry. In order to reduce future invasion risk, we suggest reviewing inter-island quarantine measures and continuing community education efforts to reduce human-mediated propagule pressure.Resumen Las especies introducidas representan la mayor amenaza a la biodiversidad terrestre única de las Islas Galápagos. En este estudio, evaluamos la condición actual de invasión de plantas en Galápagos, predecimos la posibilidad de nuevas naturalizaciones e invasiones de la flora introducida existente, y sugerimos maneras para limitar invasiones en el futuro. Ha habido un incremento de 1.46 veces en total de biodiversidad de plantas en Galápagos atribuido a las naturalizaciones de plantas introducidas; este patrón se refleja en otras islas del mundo. Han sido reportadas 870 especies de plantas introducidas en el archipiélago. De las especies evaluadas, 34% están naturalizadas; dentro de este grupo están las especies invasoras (16% de las evaluadas) y las transformadores (3.3% de las evaluadas). Como se esperaba, se muestra que las especies naturalizadas han estado presentes en el archipiélago más tiempo que las especies no-naturalizadas. También encontramos que el incremento del número de individuos de las especies introducidas sembradas por los seres humanos está directamente relacionado con el tamaño de la población humana y con la antigüedad de las propiedades privadas. Este hecho, en combinación con la introducción relativamente reciente de la mayoría de las especies, nos lleva a la conclusión de que Galápagos está en las primeras etapas de invasión. Predecimos que más de las especies introducidas ya presentes van a encontrar una oportunidad para naturalizarse a medida que ...

Native biodiversity on the Galapagos Archipelago is severely threatened by invasive alien species. On Santa Cruz Island, the abundance of introduced plant species is low in the arid lowlands of the Galapagos National Park, but increases with elevation into unprotected humid highlands. Two common alien plant species, guava (Psidium guajava) and passion fruit (Passiflora edulis) occur at higher elevations yet their seeds are dispersed into the lowlands by migrating Galapagos tortoises (Chelonoidis spp.). Tortoises transport large quantities of seeds over long distances into environments in which they have little or no chance of germination and survival under current climate conditions. However, climate change is projected to modify environmental conditions on Galapagos with unknown consequences for the distribution of native and introduced biodiversity. We quantified seed dispersal of guava and passion fruit in tortoise dung piles and the distribution of adult plants along two elevation gradients on Santa Cruz to assess current levels of ‘wasted’ seed dispersal. We computed species distribution models for both taxa under current and predicted future climate conditions. Assuming that tortoise migratory behaviour continues, current levels of “wasted” seed dispersal in lowlands were projected to decline dramatically in the future for guava but not for passion fruit. Tortoises will facilitate rapid range expansion for guava into lowland areas within the Galapagos National Park where this species is currently absent. Coupled with putative reduction in arid habitat for native species caused by climate change, tortoise driven guava invasion will pose a serious threat to local plant communities.