Rui B. Elias

Gap dynamics and regeneration strategies in Juniperus-Laurus forests of the Azores Islands

We asked the following questions regarding gap dynamics and regeneration strategies in Juniperus-Laurus forests: How important are gaps for the maintenance of tree diversity? What are the regeneration strategies of the tree species? Thirty canopy openings were randomly selected in the forest and in each the expanded gap area was delimited. Inside expanded gaps the distinction was made between gap and transition zone. In the 30 expanded gaps a plot, enclosing the gap and transition zone, was placed. In order to evaluate the differences in regeneration and size structure of tree species between forest and expanded gaps, 30 control plots were also delimited in the forest, near each expanded gap. In the 60 plots the number of seedlings, saplings, basal sprouts and adults of tree species were registered. Canopy height and width of adult individuals were also measured. The areas of the 30 gaps and expanded gaps were measured and the gap-maker identified. Juniperus-Laurus forests have a gap dynamic associated with small scale disturbances that cause the death, on average, of two trees, mainly of Juniperus brevifolia. Gap and expanded gap average dimensions are 8 and 25 m2, respectively. Gaps are of major importance for the maintenance of tree diversity since they are fundamental for the regeneration of all species, with the exception of Ilex azorica. Three types of regeneration behaviour and five regeneration strategies were identified: (1) Juniperus brevifolia and Erica azorica are pioneer species that regenerate in gaps from seedlings recruited after gap formation. However, Juniperus brevifolia is a pioneer persistent species capable of maintaining it self in the forest due to a high longevity and biomass; (2) Laurus azorica and Frangula azorica are primary species that regenerate in gaps from seedlings or saplings recruited before gap formation but Laurus azorica is able to maintain it self in the forest through asexual regeneration thus being considered a primary persistent species; (3) Ilex azorica is a mature species that regenerates in the forest.

Genetic Variability and Differentiation Among Populations of the Azorean Endemic Gymnosperm Juniperus brevifolia: Baseline Information for a Conservation and Restoration Perspective

The Azorean endemic gymnosperm Juniperus brevifolia (Seub.) Antoine is a top priority species for conservation in Macaronesia, based on its ecological significance in natural plant communities. To evaluate genetic variability and differentiation among J. brevifolia populations from the Azorean archipelago, we studied 15 ISSR and 15 RAPD markers in 178 individuals from 18 populations. The average number of polymorphic bands per population was 65 for both ISSR and RAPD. The majority of genetic variability was found within populations and among populations within islands, and this partitioning of variability was confirmed by AMOVA. The large majority of population pairwise FST values were above 0.3 and below 0.6. The degree of population genetic differentiation in J. brevifolia was relatively high compared with other species, including Juniperus spp. The genetic differentiation among populations suggests that provenance should be considered when formulating augmentation or reintroduction strategies.

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.

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.

Assessing the efficiency of protected areas to represent biodiversity: a small island case study

Protected areas (PAs) have been selected using either subjective or objective criteria applied to an extremely limited subset of biodiversity. Improved availability of species distribution data, better statistical tools to predict species distributions and algorithms to optimize spatial conservation planning allow many impediments to be overcome, particularly on small islands. This study analyses whether 219 species are adequately protected by PAs on Pico Island (the Azores, Portugal), and if they are as efficient as possible, maximizing species protection while minimizing costs. We performed distribution modelling of species’ potential distributions, proposed individual conservation targets (considering the context of each species in the archipelago and their current conservation status) to determine the efficiency of current PAs in meeting such targets and identify alternative or complementary areas relevant for conservation. Results showed that current PAs do not cover all taxa, leaving out important areas for conservation. We demonstrate that by using optimization algorithms it is possible to include most species groups in spatial conservation planning in the Azores with the current resources. With increasing availability of data and methods, this approach could be readily extended to other islands and regions with high endemism levels.

Global Island Monitoring Scheme (GIMS): a proposal for the long-term coordinated survey and monitoring of native island forest biota

Islands harbour evolutionary and ecologically unique biota, which are currently disproportionately threatened by a multitude of anthropogenic factors, including habitat loss, invasive species and climate change. Native forests on oceanic islands are important refugia for endemic species, many of which are rare and highly threatened. Long-term monitoring schemes for those biota and ecosystems are urgently needed: (i) to provide quantitative baselines for detecting changes within island ecosystems, (ii) to evaluate the effectiveness of conservation and management actions, and (iii) to identify general ecological patterns and processes using multiple island systems as repeated ‘natural experiments’. In this contribution, we call for a Global Island Monitoring Scheme (GIMS) for monitoring the remaining native island forests, using bryophytes, vascular plants, selected groups of arthropods and vertebrates as model taxa. As a basis for the GIMS, we also present new, optimized monitoring protocols for bryophytes and arthropods that were developed based on former standardized inventory protocols. Effective inventorying and monitoring of native island forests will require: (i) permanent plots covering diverse ecological gradients (e.g. elevation, age of terrain, anthropogenic disturbance); (ii) a multiple-taxa approach that is based on standardized and replicable protocols; (iii) a common set of indicator taxa and community properties that are indicative of native island forests’ welfare, building on, and harmonized with existing sampling and monitoring efforts; (iv) capacity building and training of local researchers, collaboration and continuous dialogue with local stakeholders; and (v) long-term commitment by funding agencies to maintain a global network of native island forest monitoring plots.

Comparison of -Square, Point Centered Quarter, and -Tree Sampling Methods in Pittosporum undulatum Invaded Woodlands

Tree density is an important parameter affecting ecosystems functions and management decisions, while tree distribution patterns affect sampling design. Pittosporum undulatum stands in the Azores are being targeted with a biomass valorization program, for which efficient tree density estimators are required. We compared -Square sampling, Point Centered Quarter Method (PCQM), and -tree sampling with benchmark quadrat (QD) sampling in six 900 m2 plots established at P. undulatum stands in Sao Miguel Island. A total of 15 estimators were tested using a data resampling approach. The estimated density range (344–5056 trees/ha) was found to agree with previous studies using PCQM only. Although with a tendency to underestimate tree density (in comparison with QD), overall, -Square sampling appeared to be the most accurate and precise method, followed by PCQM. Tree distribution pattern was found to be slightly aggregated in 4 of the 6 stands. Considering (1) the low level of bias and high precision, (2) the consistency among three estimators, (3) the possibility of use with aggregated patterns, and (4) the possibility of obtaining a larger number of independent tree parameter estimates, we recommend the use of -Square sampling in P. undulatum stands within the framework of a biomass valorization program.

Development of Allometric Equations for Estimating Above-Ground Biomass of Woody Plant Invaders: The Case of Pittosporum undulatum in the Azores Archipelago

The use of biomass for energy production has shown a growing interest in recent years. Pittosporum undulatum is a widespread invasive tree in the Azores archipelago with a considerable potential as an energy resource. Sustainable use of forest resources demands accurate and precise estimation of standing biomass but for the Azores only a few studies exist devoted to estimation of above-ground biomass (AGB). Here we used published allometric equations and developed new models for the estimation of P. undulatum total AGB in the exotic woodlands of the Azores. A total of 470 trees were sampled at 11 P. undulatum stands in Sao Miguel Island, and their total biomass and several dendrometric traits were recorded in the field. Several dendrometric variables were used as predictors for biomass: diameter at breast height, D; tree height,H; basal area, BA; canopy height, CH; canopy diameter, CD; canopy biovolume, BV; and number of branches at breast height, NB. For model comparison and evaluation R \(^{2}\), RMSE and AIC were used. The equations comprised models fitted to log-transformed data. There were significant and strong linear relationships between AGB and predictors BA and H. The considerable variability in P. undulatum stands, in terms of tree density and structure, should be considered when estimating biomass with allometric equations.

Modelling Native and Invasive Woody Species: A Comparison of ENFA and MaxEnt Applied to the Azorean Forest

Species distribution models are algorithmic tools that relate the distribution and occurrence of a species to the environmental characteristics of the location from where it has been recorded. Those models, also known as ecological niche models, have emerged as an effective tool in spatial ecology, conservation and land management. The Ecological Niche Factor Analysis (ENFA) is one of the common modelling approaches that are suitable for predicting potential distributions, based on presences only, providing an ecological interpretation based on marginality and specialization. In Maximum Entropy Modelling (MaxEnt) the relative entropy is minimized between the two probability densities defined in the covariate space i.e. estimated from presence data or from landscape. It focuses on relating the environmental conditions of the area where the species is present to the environmental conditions across the area of interest. ENFA has been successfully used in the Azores to model the potential distribution of indigenous and non-indigenous trees. In this paper we use distribution data from one of the most important woody plant invaders in the Azores, Pittosporum undulatum, to compare both modelling approaches. We also test both methods when using the selected environmental variables to predict the distribution in other islands and for other species (Acacia melanoxylon and Morella faya). In general, the two methodologies derived similar predictions. However, our results suggest that the set of environmental variables selected to model the distribution of a species in one particular island will probably have to be adjusted to fit other regions and species.