Beatriz Rumeu

The colonization history of Juniperus brevifolia (Cupressaceae) in the Azores Islands.

Background A central aim of island biogeography is to understand the colonization history of insular species using current distributions, fossil records and genetic diversity. Here, we analyze five plastid DNA regions of the endangered Juniperus brevifolia, which is endemic to the Azores archipelago. Methodology/Principal Findings The phylogeny of the section Juniperus and the phylogeographic analyses of J. brevifolia based on the coalescence theory of allele (plastid) diversity suggest that: (1) a single introduction event likely occurred from Europe; (2) genetic diversification and inter-island dispersal postdated the emergence of the oldest island (Santa Maria, 8.12 Ma); (3) the genetic differentiation found in populations on the islands with higher age and smaller distance to the continent is significantly higher than that on the younger, more remote ones; (4) the high number of haplotypes observed (16), and the widespread distribution of the most frequent and ancestral ones across the archipelago, are indicating early diversification, demographic expansion, and recurrent dispersal. In contrast, restriction of six of the seven derived haplotypes to single islands is construed as reflecting significant isolation time prior to colonization. Conclusions/Significance Our phylogeographic reconstruction points to the sequence of island emergence as the key factor to explain the distribution of plastid DNA variation. The reproductive traits of this juniper species (anemophily, ornithochory, multi-seeded cones), together with its broad ecological range, appear to be largely responsible for recurrent inter-island colonization of ancestral haplotypes. In contrast, certain delay in colonization of new haplotypes may reflect intraspecific habitat competition on islands where this juniper was already present.

Temporal and Spatial Variation in the Diet of the Endemic Lizard Gallotia galloti in an Insular Mediterranean Scrubland

Abstract We analyzed 621 lizard fecal pellets to assess the diet of the endemic lizard Gallotia galloti (Lacertidae) throughout one year in the thermophilous scrubland, the most threatened habitat in the Canary Islands. Indicating the importance of frugivory, 98.1% of pellets contained seeds from fleshy-fruited plant species (8,028 seeds in total), and the fruit volume reached 47.5%. The Canarian endemic plants Rhamnus crenulata and Canarina canariensis were most important in frequency of occurrence and number of seeds found in pellets, respectively. Lizards were more frugivorous during the summer (63.0% of volume), and seasonal variation was associated with temporal changes in availability of ripe fleshy fruits. We detected microspatial differences in the consumption of fruits and plant material. Significant correlations were found between plant cover of each species and their respective consumption by lizards. Lizards consumed invertebrates throughout the year; Formicidae, Hemiptera, Coleoptera, and Hymenoptera were the most frequently consumed prey (76.5% of the animal prey items). Our results indicate that G. galloti is one of the most frugivorous lacertids studied. The high number of undamaged seeds removed by these lizards also implies an important role in the seed dispersal processes of many fleshy-fruited plant species from this threatened habitat.

Differential seed dispersal systems of endemic junipers in two oceanic Macaronesian archipelagos: the influence of biogeographic and biological characteristics

This article evaluates the seed dispersal systems of two congeneric and endemic fleshy-fruited plants in the context of two relatively close oceanic archipelagos. For this purpose, representative populations of the endangered junipers Juniperus cedrus in the Canary Islands and Madeira, and Juniperus brevifolia in the Azores were studied. Despite both species sharing the same biogeographic region, we set out to test whether different conditions of the islands and biological characteristics of each juniper species determine the distinctive guilds of seed dispersers involved. We assessed the quantitative and qualitative role of the potential frugivores, showing that the wintering Turdus torquatus and the native Turdus merula were the main seed dispersers for J. cedrus and J. brevifolia, respectively (Frequency of occurrence: 74.9%, 80.2%; germination increase with respect to controls: 11.6%, 15.5%; for J. cedrus and J. brevifolia, respectively). The endemic lizard Gallotia galloti was quantitatively outstanding as seed disperser of J. cedrus, although its qualitative effect does not appear to be beneficial. The introduced rabbit Oryctolagus cuniculus acts as a disruptor in both natural seed dispersal systems, as inferred from the high percentage of damaged seeds found in their droppings. Our results indicate that J. cedrus and J. brevifolia are primarily adapted to ornithochory processes, T. torquatus and T. merula being their respective legitimate long-distance dispersers. Although these birds should be playing a key role in the connectivity of fragmented populations, the dependence of J. cedrus on a migrant bird involves a notable fragility of the system.

Contrasting phenology and female cone characteristics of the two Macaronesian island endemic cedars (Juniperus cedrus and J. brevifolia)

Phenology and female cone characteristics of the two endemic cedars (Juniperus cedrus and J. brevifolia) from the Macaronesian islands were studied. Despite their closely taxonomic affinity and their evolution under insular conditions, different trends were recorded. Mature J. cedrus female cones were present throughout the year, while those from J. brevifolia were only present in summer and autumn. J. cedrus female cone size was significantly larger than that of J. brevifolia, a trend consistent with the presence of larger vertebrates (lizards and birds) in the Canary Islands. However, water content was four times higher in J. brevifolia female cones, which can be related with the higher rainfall existing in the Azores. J. cedrus has two or three seeds per cone, whereas J. brevifolia frequently had three. Seeds from J. cedrus were clearly larger and heavier, coinciding with the female cone size trend. However, tetrazolium tests revealed higher viability values in J. brevifolia. The relatively low percentage of filled seeds in J. cedrus could be a consequence of the climatic stress and limits to pollination due to fragmented populations as described for other Juniperus species. In summary, our results reveal that some environmental factors such as the harsh conditions, high population fragmentation and the dependence on large dispersers have compromised the fitness of J. cedrus in the Canary Islands.

Seasonal Diet of the Grey Heron Ardea cinerea on an Oceanic Island (Tenerife, Canary Islands): Indirect Interaction with Wild Seed Plants

Abstract. In 199 pellets analyzed a total of 7 460 prey items were counted, 96.2% of which were arthropods. Aeshnidae larvae (Odonata) made up 66.1% of the total prey items and were the main invertebrate group. Vertebrates constituted 3.8%, with reptiles and mammals being the main prey of this type (1.8% each). Despite the small size of the invertebrates, this group reached > 60% in terms of biomass. All the main prey items varied significantly among seasons. Odonata was the most important group in all seasons, reaching its maximum value in summer. In the case of vertebrates, reptiles were captured mainly in spring, mammals in winter. With regard to indirect interaction with seeds, a total of 901 seeds associated with lizard remains were found in 77 pellets, indicating that they had previously been consumed by these reptiles. External visual damage of seeds was low and only 1.1% was destroyed. No seeds germinated after the four-month germination experiment and practically all of them were unviable. In conclusion, these results indicate that Grey Heron diet on islands varies in comparison with continental zones, including an important number of invertebrates and reptiles. Furthermore, this bird acts as an opportunistic secondary seed disperser, although its ecological effect does not seem to be very significant for the dynamics of the Canary Island ecosystems.

Predicting the consequences of disperser extinction: richness matters the most when abundance is low

Summary 1. The ongoing biodiversity crisis entails the concomitant loss of species and the ecological services they provide. Global defaunation, and particularly the loss of frugivores, may negatively affect the seed dispersal of fleshy-fruited plant species, with predictable stronger impacts in simplified communities such as those on oceanic islands. However, logistical difficulties have hindered the experimental and theoretical need to disentangle the roles of species identity, richness (i.e. number of species) and abundance. Consequently, studies to date have focused exclusively on the loss of species richness leaving us largely ignorant regarding how species identity and abundance affect the loss of ecosystem functions. 2. Here, we applied a network approach to disentangle the effects of disperser abundance, richness and identity on the seed dispersal service provided by frugivores to the Galapagos plant community. 3. We found that both abundance and richness of the dispersers significantly affect the function of seed dispersal and that richness becomes increasingly important as disperser abundance declines. Extinction simulations revealed that the order of species loss has profound implications to the plant community. On one hand, abundant generalist dispersers like the Galapagos lizards, can mitigate the loss of specialized dispersers. On the other hand, specific threats affecting key dispersers can lead to the rapid collapse of the community-level dispersal services. 4. Our results suggest that the identity of the disperser species lost can have a large effect on the number of plant species dispersed, and generalist species are essential to the persistence of the community dispersal service. Both abundance and species richness of seed dispersers are key and synergistic drivers of the number of plant species dispersed. Consequently, the combined effect of their loss in degraded ecosystems can result in positive feedbacks, further accelerating the decline of the dispersal function. This article is protected by copyright. All rights reserved.

Galapagos land iguana (Conolophus subcristatus) as a seed disperser.

The role of the most common land iguana (Conolophus subcristatus) in the Galápagos Islands as an effective seed disperser is explored in this study. A total of 5705 seeds of 32 plant species were identified from 160 scats, 4545 of which (80%) appeared visually undamaged. Germination trials of 849 seeds from 29 species revealed that at least 10 species remained viable after passing through the iguanas gut, although only a small proportion of those seeds (4%) germinated. In any case, we argue that C. subcristatus exerts an important role on the 7 Galapagos islands where it occurs because of its abundance and capacity to ingest and disperse seeds at long distances. Our results strongly suggest that the Galápagos C. subcristatus plays an important role as a seed disperser of not only of native species but also some introduced plants in the Galápagos Islands.

Contribution by vertebrates to seed dispersal effectiveness in the Galápagos Islands: a community-wide approach

Seed dispersal and seedling recruitment are crucial phases in the life cycle of all spermatophyte plants. The net contribution of seed dispersers to plant establishment is known as seed dispersal effectiveness (SDE) and is defined as the product of a quantitative (number of seeds dispersed) and a qualitative (probability of recruitment) component. In Galápagos, we studied the direct contribution to SDE (number of seeds dispersed and effect on seedling emergence) provided by the five island groups of frugivores (giant tortoises, lizards, medium-sized passerine birds, small non-finch passerine birds, and finches) in the two main habitats in this archipelago: the lowland and the highland zones, and found 16 vertebrate species dispersing 58 plant species. Data on frequency of occurrence of seeds in droppings and number of seeds dispersed per unit area produced contrasting patterns of seed dispersal. Based on the former, giant tortoises and medium-sized passerines were the most important seed dispersers. However, based on the latter, small non-finch passerines were the most important dispersers, followed by finches and medium-sized passerines. The effect of disperser gut passage on seedling emergence varied greatly depending on both the disperser and the plant species. Although the contribution to SDE provided by different disperser guilds changed across plant species, medium-sized passerines (e.g., mockingbirds) provided a higher contribution to SDE than lava lizards in 10 out of 16 plant species analysed, whereas lava lizards provided a higher contribution to SDE than birds in five plant species. While both the quantitative and qualitative components addressed are important, our data suggests that the former is a better predictor of SDE in the Galápagos archipelago.

Historical isolation of the Galápagos carpenter bee (Xylocopa darwini) despite strong flight capability and ecological amplitude

Colonization across the Galápagos Islands by the carpenter bee (Xylocopa darwini) was reconstructed based on distribution of mitochondrial haplotypes (cytochrome oxidase II (COII) sequences) and haplotype lineages. A total of 12 haplotypes were found in 118 individuals of X. darwini. Distributional, phylogenetic and phylogeographic analyses suggest early colonization of most islands followed by historical isolation in two main groups: eastern and central-western islands. Evidence of recurrent inter-island colonization of haplotypes is largely lacking, despite strong flight capability and ecological amplitude of the species. Recent palaeogeographic data suggest that several of the current islands were connected in the past and thus the isolation pattern may have been even more pronounced. A contrast analysis was also carried out on 10 animal groups of the Galápagos Islands, and on haplotype colonization of seven animal and plant species from several oceanic archipelagos (the Galápagos, Azores, Canary Islands). New colonization metrics on the number of potential vs. inferred colonization events revealed that the Galápagos carpenter bee shows one of the most significant examples of geographic isolation.

Newly Discovered Seed Dispersal System of Juniperus cedrus Questions the Pristine Nature of the High Elevation Scrub of El Teide (Tenerife, Canary Islands)

Abstract As a working hypothesis, we examined evidence for the former presence of a climacic woodland of Juniperus cedrus above the pine forest in the high elevation area of Tenerife (Canary Islands), which would indicate that the current dominant vegetation (endemic Spartocytisus supranubius scrub) may not be pristine. The main causes of the great regression of this woodland were caused by human activities (timber harvesting, herbivory by goats, and fires). The main support for this hypothesis is the survival of a presumably relict seed dispersal system of the endangered endemic J. cedrus, which relies mainly on the wintering thrush Turdus torquatus. The fact that genetic factors are directly involved in the control of bird migration routes strongly supports the idea that this interaction could be remnant of an older system, probably more widespread in the past. To test this hypothesis, we propose that a paleoecological approach could reconstruct the vegetation dynamics in the Teide National Park (Tenerife) and the past presence of this seed disperser migratory thrush. The analysis of plant microfossils in sediments (e.g., pollen, spores, phytoliths, coprolites, and charcoal) would allow us to evaluate whether the current vegetation is the same as that which naturally existed in the past, and assess the impact of the anthropogenic and natural factors to which it has been subjected during history. The results of these analyses will be useful for future management policies and practices aimed at restoring the pristine landscape and biotic interactions of the Teide National Park. To our knowledge, the case presented in this contribution, based on the high dependence of the seed dispersal of an endemic tree (J. cedrus) on a migratory bird, is the only reported in the context of oceanic islands.