Systematics,palaeoecology and palaeobiogeography of the Neogene fossil sharks from the Azores (Northeast Atlantic)

The Azores Archipelago is a group of isolated islands located in the North Atlantic Ocean. One of these oceanic islands – Santa Maria – exhibits marine fossiliferous sediments of late Miocene/early Pliocene and also of Pleistocene age. Recent research provided new selachian fossil material, with three new records (Carcharias acutissima, Megaselachus megalodon, and Carcharhinus cf. leucas) increasing the number of fossil sharks reported from the Azores (Santa Maria Island) to seven species (Notorynchus primigenius, C. acutissima, Cosmopolitodus hastalis, Paratodus benedenii, Isurus oxyrinchus, M. megalodon, and C. cf. leucas). So far, no teeth of batoids or small sharks have been found despite the screen-washing of several sediment samples from Santa Maria. The Azorean Mio-Pliocene selachian fauna clearly differs from those described from sediments deposited on continental shelves, in which batoids and small benthic sharks (e.g., scyliorhinids) are usually well represented. During the late Miocene/early Pliocene, subtropical to warm-temperate seas were prevalent in the area of the Azores, as deduced from palaeontological, geological and isotopic data, all indicating a warmer climate than in the present.     

Late Miocene marine ostracods from Santa Maria island,Azores (NE Atlantic): Systematics,palaeoecology and palaeobiogeography

The nine oceanic islands that comprise the Azores archipelago are located in the middle of the northern Atlantic Ocean. In this isolated archipelago, there is a rich fossil record in one of the islands, Santa Maria. In this island, samples were collected in the Upper Miocene composite section of Malbusca outcrop, located in the southern shore of the island, and the fossil marine Ostracoda were studied. This work represents the first report of fossil ostracods from the Azores archipelago. Thirteen species were found, representing seven families and 12 genera (Xestoleberis, Loxoconcha, Callistocythere, Leptocythere, Dameriacella, Aurila, Heliocythere, Pachycaudites, Neonesidea, Cyamocytheridea, ?Quadracythere and Paracypris). Among the identified species, one new species, Leptocythere azorica n. sp., is described. Loxoconcha (two species) was the most diversified genus. The collected species are mainly ornamented and typical of warm waters and epi-neritic habitats (～ 10–50 m of depth).     

Phylogeography and molecular phylogeny of Macaronesian island Tarphius (Coleoptera: Zopheridae): why are there so few species in the Azores?

Aim We used a phylogenetic framework to examine island colonization and predictions pertaining to differentiation within Macaronesian Tarphius (Insecta, Coleoptera, Zopheridae), and explain the paucity of endemics in the Azores compared with other Macaronesian archipelagos. Specifically, we test whether low diversity in the Azores could be due to recent colonization (phylogenetic lineage youth), cryptic speciation (distinct phylogenetic entities within species) or the young geological age of the archipelago. Location Macaronesian archipelagos (Azores, Madeira and the Canary Islands), northern Portugal and Morocco. Methods Phylogenetic analyses of mitochondrial and nuclear genes of Tarphius beetles of the Azores, other Macaronesian islands and neighbouring continental areas were used to investigate the origin of island biodiversity and to compare patterns of colonization and differentiation. A comparative nucleotide substitution rate test was used to select the appropriate substitution rate to infer clade divergence times. Results Madeiran and Canarian Tarphius species were found to be more closely related to each other, while Azorean taxa grouped separately. Azorean taxa showed concordance between species and phylogenetic clades, except for species that occur on multiple islands, which segregated by island of origin. Divergence time estimates revealed that Azorean Tarphius are an old group and that the most recent intra‐island speciation event on Santa Maria, the oldest island, occurred between 3.7 and 6.1 Ma. Main conclusions Our phylogenetic approach provides new evidence to understand the impoverishment of Azorean endemics: (1) Tarphius have had a long evolutionary history within the Azores, which does not support the hypothesis of fewer radiation events due to recent colonization; (2) the current taxonomy of Azorean Tarphius does not reflect common ancestry and cryptic speciation is responsible for the underestimation of endemics; (3) intra‐island differentiation in the Azores was found only in the oldest island, supporting the idea that young geological age of the archipelago limits the number of endemics; and (4) the lack of evidence for recent intra‐island diversification in Santa Maria could also explain the paucity of Azorean endemics. Phylogenetic reconstructions of other species‐rich taxa that occur on multiple Macaronesian archipelagos will reveal whether our conclusions are taxon specific, or of a more general nature.     

Genetic structure of house mouse (Mus musculus Linnaeus 1758) populations in the Atlantic archipelago of the Azores: colonization and dispersal

We analyzed the genetic structure and relationships of house mouse (Mus musculus) populations in the remote Atlantic archipelago of the Azores using nuclear sequences and microsatellites. We typed Btk and Zfy2 to confirm that the subspecies Mus musculus domesticus was the predominant genome in the archipelago. Nineteen microsatellite loci (one per autosome) were typed in a total of 380 individuals from all nine Azorean islands, the neighbouring Madeiran archipelago (Madeira and Porto Santo islands), and mainland Portugal. Levels of heterozygosity were high on the islands, arguing against population bottlenecking. The Azorean house mouse populations were differentiated from the Portuguese and Madeiran populations and no evidence of recent migration between the three was obtained. Within the Azores, the Eastern, Western, and Central island groups tended to act as separate genetic units for house mice, with some exceptions. In particular, there was evidence of recent migration events among islands of the Central island group, whose populations were relatively undifferentiated. Santa Maria had genetically distinctive mice, which may relate to its colonization history. © 2013 The Linnean Society of London     

Environmental variability of Macaronichnus ichnofabrics in Eocene tidal‐embayment deposits of southern Patagonia,Argentina

The Middle Eocene Man Aike Formation of southern Patagonia, Argentina, offers the opportunity to study the palaeoecological and palaeoenvironmental significance of Macaronichnus ichnofabrics in a tide‐dominated, coastal embayment compound‐dune complex. Four recurrent Macaronichnus ichnofabrics are recognized: Macaronichnus IF‐1 characterized by Macaronichnus segregatis segregatis; Macaronichnus IF‐2 with M. segregatis segregatis, Ophiomorpha nodosa, Skolithos linearis and rare Palaeophycus tubularis; Macaronichnus IF‐3 typified by M. segregatis segregatis, O. nodosa, Asterosoma radiciforme and rare S. linearis; and Macaronichnus IF‐4 consisting of M. segregatis segregatis, Nereites missouriensis, A. radiciforme, O. nodosa, Rosselia socialis, S. linearis, P. tubularis, Planolites isp. and Chondrites isp. Variations in these ichnofabrics are linked to changes in local hydraulic conditions in the upper and lower trough areas of the compound dunes. In high‐energy, shallow and marginal marine environments, the Macaronichnus tracemaker is commonly the first to colonize the shifting, sandy substrates. Overprinting by other ichnoguilds occurs only after the physiochemical conditions became favourable and the colonization window length increases.     

ORIGINAL ARTICLE: Mass extinctions in the Azores during the last glaciation: fact or myth?

Aim The influence of the last glaciation on the shallow‐water marine malacofauna of the Azores Islands is reviewed. We test, for this fauna, the ‘Pleistocene temperature theory’ of J.C. Briggs, which hypothesizes that a (supposed) lack of endemics in the older (Azorean endemic) fauna resulted from extinctions caused by a severe drop in sea surface temperatures during the Pleistocene. Location Santa Maria Island, Azores, Portugal. Methods We compare the fossil mollusc fauna of Prainha, Praia do Calhau and Lagoinhas Pleistocene outcrops with the recent mollusc fauna of the Azores Islands. We dated the fossil fauna, using shells of Patella aspera Röding, 1798, by standard U/Th methodology at the GEOTOP laboratory (Université du Québec à Montreal, Canada). Results Dating of the shells of P. aspera indicates that the deposition of the lower unit of the Prainha outcrop corresponded to Marine Oxygen Isotope Substage 5e (MISS 5e). Not a single endemic Azorean species of mollusc that is present in the Pleistocene fossil record has since become extinct, and we found no signs of ‘mass extinctions’ in the littoral marine molluscs of the Azores. The only species that were extirpated from these islands were thermophilic molluscs and littoral bivalves living in fine sand. Main conclusions Our results do not support Briggs’‘Pleistocene temperature theory’. Nor did we find evidence supporting the hypothesis that most of the marine organisms now present in the Azores recolonized the islands after the last glacial maximum.     

Song dialects on the Atlantic islands: goldcrests of the Azores ( Regulus regulus azoricus, R. r. sanctae-mariae, R. r. inermis)

Territorial songs in island populations of songbirds are often highly divergent from those of neighbouring continental relatives. This is shown for the three goldcrest subspecies (Regulus regulus azoricus, R. r. sanctae-mariae and R. r. inermis) endemic on six islands of the Azorean archipelago. All investigated populations display a high intra- and inter-individual acoustic variation. On each island, up to six different song types have been found; and a single male sings up to three types. In contrast, all northwestern European populations of R. r. regulus and R. r. anglorum share only a single song type. In playback experiments, none of 18 tested dialect songs of Azorean goldcrests evoked notable territorial reaction in German and Czech goldcrest males (ssp. regulus). Two differing dialect groups of the goldcrest can be distinguished on the Azores. Populations of the eastern islands, São Miguel and Santa Maria, share common song types which are not found on the islands of the central and western groups. Dialect repertoires on the westernmost islands, Flores, Faial and Pico, are dominated by a different song type. In the geologically younger western crater of São Miguel, both western and eastern song types coexist. Acoustic similarities to a population from neighbouring Terceira suggest the western part of São Miguel as the origin for the westward expansion of R. regulus on the Azores.     

Dynamics of Whale Shark Occurrence at Their Fringe Oceanic Habitat

Studies have shown that the whale shark (Rhincodon typus), a vulnerable large filter feeder, seasonally aggregates at highly productive coastal sites and that individuals can perform large, trans-boundary migrations to reach these locations. Yet, the whereabouts of the whale shark when absent from these sites and the potential oceanographic and biological drivers involved in shaping their present and future habitat use, including that located at the fringes of their suitable oceanic habitat, are largely unknown. We analysed a 16-year (1998–2013) observer dataset from the pole-and-line tuna fishery across the Azores (mid-North Atlantic) and used GAM models to investigate the spatial and temporal patterns of whale shark occurrence in relation to oceanographic features. Across this period, the whale shark became a regular summer visitor to the archipelago after a sharp increase in sighting frequency seen in 2008. We found that SST helps predicting their occurrence in the region associated to the position of the seasonal 22°C isotherm, showing that the Azores are at a thermal boundary for this species and providing an explanation for the post 2007 increase. Within the region, whale shark detections were also higher in areas of increased bathymetric slope and closer to the seamounts, coinciding with higher chl-a biomass, a behaviour most probably associated to increased feeding opportunities. They also showed a tendency to be clustered around the southernmost island of Santa Maria. This study shows that the region integrates the oceanic habitat of adult whale shark and suggests that an increase in its relative importance for the Atlantic population might be expected in face of climate change.     

Tracking macroalgae introductions in North Atlantic oceanic islands

The Azores archipelago was selected as a case study since there are few studies on macroalgae introduction in oceanic islands. While at a global scale, around 3 % of macroalgae are considered non-indigenous; in the remote oceanic islands of the Azores, over 6 % of the marine algal flora is non-indigenous. The taxa distribution pattern of non-indigenous species in the Azores is significantly different from the distribution pattern in the globe. The most representative group was Rhodophyta species, being 84 % of the total non-indigenous macroalgae, mainly introduced via maritime traffic. This study highlights the vulnerability of remote islands to the introduction of macroalgae and the need to develop further studies on other archipelagos to understand whether the observed vulnerability is generally characteristic of oceanic islands. The development of local monitoring and mitigation programs and the necessity of regulatory and preventive measures for the maritime traffic vector are strongly suggested.     

Time, area and isolation: factors driving the diversification of Azorean arthropods

Aim R. J. Whittaker et al. recently proposed a ‘general dynamic model of oceanic island biogeography’ (GDM), providing a general explanation of island biodiversity patterns by relating fundamental biogeographical processes – speciation, immigration, extinction – to area (A) and time (T; maximum island geological age). We adapt their model, which predicts a positive relationship with area combined with a humped relationship to time (designated the ATT² model), to study the factors promoting diversification on the Azores for several arthropod groups. Location The Azorean archipelago (North Atlantic; 37–40° N, 25–31° W). Methods We use the number of single‐island endemics (SIEs) as a measure of diversification, to evaluate four different predictions for the variation in SIEs between different islands, derived from the GDM theory and our knowledge of the fauna and history of the Azores. We calculated the number of SIEs for seven out of the nine Azorean islands and six groups of species (all arthropods, beetles, cavernicolous and non‐cavernicolous species, and taxa with high and low dispersal abilities). Several variables accounting for island characteristics (area, geological age, habitat diversity and isolation) and generalized linear models were used to evaluate the reliability of each prediction. Results A linear and positive relationship between SIEs and an AT (area + time) model was the most parsimonious explanation for overall arthropod diversification. However, cavernicolous species showed the opposite pattern (more SIEs inhabiting the youngest islands). Also, isolation was an important predictor of diversification for all groups except for the species with high dispersal ability; while the former were negatively related to the distance from the main source of colonizing lineages (Santa Maria island in most cases), the latter were related to area. Dispersal ability was also a key factor affecting the diversification of most groups of species. Main conclusions In general, the diversification of Azorean arthropods is affected by age, area and isolation. However, different groups are affected by these factors in different ways, showing radically different patterns. Although the ATT² model fails to predict the diversification pattern of several groups, it provides a framework for integrating these deviations into a general theory. Further improvements of the GDM theory need to take into account the particular traits of each group and the role of isolation in shaping island diversity.     

Algae-based biotopes of the Azores (Portugal): spatial and seasonal variation

The increasing importance of coastal management created the need for a systematic classification and characterization of marine communities. Accurate quantitative methodologies for rocky shore algae-based biotope definition, were developed and tested on the Islands of São Miguel and Santa Maria (Azores). Shores of both islands were surveyed, covering all rocky substrate types. Biotopes were defined by assessing the associated habitat and species characteristics, using ANOSIM and SIMPER analysis, respectively. A total of ten biotopes were identified. Generally both islands’ biotopes are characterized by the same taxa/ecological categories, in summer and in winter. However, association between these taxa/ecological categories and the shore height at which they occur differs geographically and temporally. There is a generalized gradual succession of taxa/ecological categories from upper intertidal down to deepest subtidal, although geographical differences occur. Diversity is highest at the land–water interface and decreases towards both extremes (upper intertidal and deepest subtidal level). The strongest evidence of seasonal variation occurs at the upper intertidal. The methodology used proves to be effective in broad scale shoreline assessment of biological communities in warm-temperate coastal marine environments, and thus suitable for the purpose it was developed for. As a consequence it should be applied to the remaining islands of the Azorean archipelago as well as to other macaronesian islands, e.g. Madeira and the Canaries.     

Invertébrés Sans Frontières: Large Scales of Connectivity of Selected Freshwater Species among Caribbean Islands

The freshwater fauna (crustaceans, molluscs, fish) of many tropical islands in the Caribbean and Pacific share an amphidromous life‐cycle, meaning their larvae need to develop in saline conditions before returning to freshwater as juveniles. This community dominates the freshwaters of much of the tropics, but is poorly known and at risk from development, in particular dam construction. Amphidromy can theoretically lead to dispersal between different freshwater areas, even to distant oceanic islands, via the sea. The extent and scale of this presumed dispersal, however, is largely unknown in the Caribbean. Recent genetic work in Puerto Rico has shown that many freshwater species have little or no population structure among different river catchments, implying high levels of connectivity within an island, whereas between‐island structure is unknown. We used genetic techniques to infer the geographic scales of population structure of amphidromous invertebrates (a gastropod and a number of crustacean species) between distant parts of the Caribbean, in particular Puerto Rico, Panama and Trinidad. We found virtually no geographic population structure across over 2000 km of open sea for these freshwater species. This implies that they are indeed moving between islands in sea currents as larvae, meaning that continued recruitment requires a continuum of healthy habitat from the freshwater to marine environment. We further discuss the role of amphidromy and suggest its ecological and biogeographic role may be more important than previously presumed.     

Genetic differentiation reflects geological history in the Azorean land snail,Leptaxis azorica

The land snail Leptaxis azorica, endemic on the Azores, was subjected to an electrophoretic (allozymes) and morphometric (genital tract) analysis. Genetic distances suggest the presence of four distinct lineages and are compatible with colonisation proceeding from the eastern, older islands (Santa Maria and S?o Miguel) to the west (Flores and Corvo). On S?o Miguel, genetic and morphometric differentiation is concordant with the separate colonisation of two islands that gave rise to the current island 50,000 years ago. The maximum time available for differentiation in isolation (0.55 million years) suggests a high rate of allozyme change between the two lineages on S?o Miguel. This may be related to population isolation and bottlenecks caused by human and volcanic activity on S?o Miguel in relatively recent times. This is more prominent in the eastern region where populations are also characterised by reduced genetic variation (loss of alleles and heterozygosity) compared to populations elsewhere.     

Drivers of diversity in Macaronesian spiders and the role of species extinctions

Aim To identify the biogeographical factors underlying spider species richness in the Macaronesian region and assess the importance of species extinctions in shaping the current diversity. Location The European archipelagos of Macaronesia with an emphasis on the Azores and Canary Islands. Methods Seven variables were tested as predictors of single‐island endemics (SIE), archipelago endemics and indigenous spider species richness in the Azores, Canary Islands and Macaronesia as a whole: island area; geological age; maximum elevation; distance from mainland; distance from the closest island; distance from an older island; and natural forest area remaining per island – a measure of deforestation (the latter only in the Azores). Different mathematical formulations of the general dynamic model of oceanic island biogeography (GDM) were also tested. Results Island area and the proportion of remaining natural forest were the best predictors of species richness in the Azores. In the Canary Islands, area alone did not explain the richness of spiders. However, a hump‐shaped relationship between richness and time was apparent in these islands. The island richness in Macaronesia was correlated with island area, geological age, maximum elevation and distance to mainland. Main conclusions In Macaronesia as a whole, area, island age, the large distance that separates the Azores from the mainland, and the recent disappearance of native habitats with subsequent unrecorded extinctions seem to be the most probable explanations for the current observed richness. In the Canary Islands, the GDM model is strongly supported by many genera that radiated early, reached a peak at intermediate island ages, and have gone extinct on older, eroded islands. In the Azores, the unrecorded extinctions of many species in the oldest, most disturbed islands seem to be one of the main drivers of the current richness patterns. Spiders, the most important terrestrial predators on these islands, may be acting as early indicators for the future disappearance of other insular taxa.     

Effect of island geological age on the arthropod species richness of Azorean pastures

Species richness of six pasture arthropod assemblages (total arthropod species, total herbivore species, sucking and chewing herbivores, total predatory species and spiders) were regressed against several geographical variables (area, distance from the nearest mainland, maximum elevation and geological age of the islands) of three Azorean islands (S. Maria, Terceira and Pico). The species were sampled by the fixed-quadrat size sampling method and the results obtained are consistent with the geological age hypothesis, i.e. the species richness of the six indigenous arthropod assemblages increases with the geological age of the islands, both at local and regional scales. Higher values of indigenous and endemic species richness were consistendy found on the older island (S. Maria), and the lowest values on the most recent island (Pico). Moreover, when considering the age of Faial (an older island probably once connected with Pico) as a estimate of the age of Pico, correlations between species richness and island age were improved, thereby strengthening the relationship. The older island (S. Maria) has more specialized herbivores and a greater proportion of herbivores in relation to predatory arthropods. Ecological and biogeographical studies in the Azores should take into account the effects of the time each island has been available for colonization and evolution.     

Towards a ‘Sea‐Level Sensitive’ dynamic model: impact of island ontogeny and glacio‐eustasy on global patterns of marine island biogeography

A synthetic model is presented to enlarge the evolutionary framework of the General Dynamic Model (GDM) and the Glacial Sensitive Model (GSM) of oceanic island biogeography from the terrestrial to the marine realm. The proposed ‘Sea‐Level Sensitive’ dynamic model (SLS) of marine island biogeography integrates historical and ecological biogeography with patterns of glacio‐eustasy, merging concepts from areas as diverse as taxonomy, biogeography, marine biology, volcanology, sedimentology, stratigraphy, palaeontology, geochronology and geomorphology. Fundamental to the SLS model is the dynamic variation of the littoral area of volcanic oceanic islands (defined as the area between the intertidal and the 50‐m isobath) in response to sea‐level oscillations driven by glacial–interglacial cycles. The following questions are considered by means of this revision: (i) what was the impact of (global) glacio‐eustatic sea‐level oscillations, particularly those of the Pleistocene glacial–interglacial episodes, on the littoral marine fauna and flora of volcanic oceanic islands? (ii) What are the main factors that explain the present littoral marine biodiversity on volcanic oceanic islands? (iii) How can differences in historical and ecological biogeography be reconciled, from a marine point of view? These questions are addressed by compiling the bathymetry of 11 Atlantic archipelagos/islands to obtain quantitative data regarding changes in the littoral area based on Pleistocene sea‐level oscillations, from 150 thousand years ago (ka) to the present. Within the framework of a model sensitive to changing sea levels, we discuss the principal factors affecting the geographical range of marine species; the relationships between modes of larval development, dispersal strategies and geographical range; the relationships between times of speciation, modes of larval development, ecological zonation and geographical range; the influence of sea‐surface temperatures and latitude on littoral marine species diversity; the effect of eustatic sea‐level changes and their impact on the littoral marine biota; island marine species–area relationships; and finally, the physical effects of island ontogeny and its associated submarine topography and marine substrate on littoral biota. Based on the SLS dynamic model, we offer a number of predictions for tropical, subtropical and temperate volcanic oceanic islands on how rates of immigration, colonization, in‐situ speciation, local disappearance, and extinction interact and affect the marine biodiversity around islands during glacials and interglacials, thus allowing future testing of the theory.     

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.     

Canopy insect herbivores in the Azorean Laurisilva forests: key host plant species in a highly generalist insect community

This article explores patterns of insect herbivore distribution in the canopy of the Laurisilva forests on seven islands in the Azores archipelago. To our knowledge, this is one of the first extensive study of this type in tree or shrub canopies of oceanic island ecosystems. One of the most frequently debated characteristics of such ecosystems is the likely prevalence of vague, ill‐defined niches due to taxonomic disharmony, which may have implications for insect‐plant interactions. For instance, an increase in ecological opportunities for generalist species is expected due to the lack of predator groups and reduced selection for chemical defence in host plants. The following two questions were addressed: 1) Are specialists species rare, and insect herbivore species randomly distributed among host plant species in the Azores? 2) Are the variances in insect herbivore species composition, frequency and richness explained by host plants or by regional island effects? We expect a proportional distribution of herbivore species between host plants, influenced by host frequency and distinct island effects; otherwise, deviation from expectation might suggest habitat preference for specific host tree crowns. Canopy beating tray samples were performed on seven islands, comprising 50 transects with 1 to 3 plant species each (10 replicates per species), giving 1320 samples from ten host species trees or shrubs in total. From a total of 129 insect herbivore species, a greater number of herbivore species was found on Juniperus brevifolia (s=65) and Erica azorica (s=53). However, the number of herbivore species per individual tree crown was higher for E. azorica than for any other host, on all islands, despite the fact that it was only the fourth more abundant plant. In addition, higher insect species richness and greater insect abundance were found on the trees of Santa Maria Island, the oldest in the archipelago. Insect species composition was strongly influenced by the presence of E. azorica, which was the only host plant with a characteristic fauna across the archipelago, whereas the fauna of other plant crowns was grouped by islands. The great insect occurrence on E. azorica reflects strong habitat fidelity, but only four species were clearly specialists. Our findings indicate a broadly generalist fauna. The simplicity of Azorean Laurisilva contributed to the understanding of insect‐plant mechanisms in canopy forest habitats.     

On the Limited Potential of Azorean Fleshy Fruits for Oceanic Dispersal

How plants arrived to originally sterile oceanic islands has puzzled naturalists for centuries. Dispersal syndromes (i.e., diaspore traits that promote dispersal by long-distance dispersal vectors), are generally considered to play a determinant role in assisting island colonization. However, the association between diaspore traits and the potential vectors by which diaspores are dispersed is not always obvious. Fleshy fruits, in particular, are considered to have evolved to promote the internal dispersal of seeds by frugivores (endozoochory), however some fleshy fruits can also float in saltwater, and thus be potentially transported by oceanic current (thalassochory). We performed saltwater floatation and viability experiments with fruits of the 14 European fleshy-fruited species that naturally colonized the Azores archipelago (North Atlantic Ocean). We show that only Corema album (a berry) and Juniperus oxycedrus (a fleshy cone) floated for as long as 60 days, the estimated minimum time needed to reach the Azores by oceanic currents. Regardless the floatation potential, exposure to saltwater largely reduced the viability of most seeds of the 14 species (46% of viability decline within 15 days and 77% within 60 days of immersion), including those of Corema album (61%) and Juniperus oxycedrus (83%). Floatability and viability trials suggest that while some fleshy-fruited species might have arrived to the Azores by oceanic currents, such would have required extreme meteorological events that could largely reduce the duration of the trip. Thus, the alternative hypothesis that fleshy-fruited species were mostly dependent on animal dispersers (endozoochory) to colonize these remote islands is reinforced.