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.     

Additive partitioning as a tool for investigating the flora diversity in oceanic archipelagos

This paper introduces the integration of additive partitioning with species—area relationships to island biogeography in order to address the question “How are the pteridophyte and spermatophyte native and endemic flora of different oceanic archipelagos partitioned across islands?”.Species richness data of all endemic species and all native species of pteridophytes and spermatophytes were obtained for the Azores, Canaries and Cape Verde in the Atlantic Ocean and Galápagos, Hawaii and Marquesas in the Pacific Ocean. Additive partitioning of species diversity was used to quantify how much of the total diversity of an oceanic archipelago flora (γ-diversity) is due to (i) the mean species richness of the flora of each island (α-diversity), (ii) the variability in species richness of the floras across islands (β_Nestedness) and (iii) the complementarity in species composition of the floras of different islands (β_Replacement). The analysis was separately performed for the native and endemic pteridophyte and spermatophyte floras.The diversity partitioning of the six archipelagos showed large differences in how the flora of each archipelago is partitioned among the α, β_Nestedness and β_Replacement components, for pteridophytes and spermatophytes and for all endemic species and all native species. The α-diversity was more important for all native species than for endemic species and more important for pteridophytes than for spermatophytes, with the Azores showing outstanding high values of α-diversity. The β_Nestedness was higher for pteridophytes than for spermatophytes and higher for endemic species than for all native species in both pteridophytes and spermatophytes. The values of β_Replacement suggested that: (i) the spermatophyte native flora is more differentiated across islands than the pteridophyte native flora and (ii) the pteridophyte endemic flora and, especially, the spermatophyte endemic flora are more differentiated across islands than the corresponding native flora. An outstanding value of β_Replacement for endemic and all native spermatophytes was found in Hawaii, confirming the biogeographical island differentiation in this archipelago.     

Pollination,biogeography and phylogeny of oceanic island bellflowers (Campanulaceae)

We studied the pollination biology of nine island Campanulaceae species: Azorina vidalii, Musschia aurea, M. wollastonii, Canarina canariensis, Campanula jacobaea, Nesocodon mauritianus, and three species of Heterochaenia. In addition, we compared C. canariensis to its two African mainland relatives C. eminii and C. abyssinica. We asked to what extent related species converge in their floral biology and pollination in related habitats, i.e. oceanic islands. Study islands were the Azores, Madeira, Canary Islands, Cape Verde, Mauritius, and Réunion. Information about phylogenetic relationships of these species and their relatives were gathered from atpB, matK, rbcL and trnL-F regions, building the most complete phylogeny of Campanulaceae to date. Six of the island bellflower species were bird-pollinated and two (A. vidalii and M. aurea) were lizard-pollinated. Insects also visited some of the species, and at least C. jacobaea had both insect- and self-pollination. Several morphological traits were interpreted as adaptations to bird and lizard pollination, e.g. all had a robust flower morphology and, in addition, bird-pollinated species were scentless, whereas lizard-pollinated species had a weak scent. These examples of vertebrate pollination evolved independently on each island or archipelago. We discuss if these pollination systems have an island or mainland origin and when they may have evolved, and finally, we attempt to reconstruct the pollinator-interaction history of each species.     

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.     

Genetic diversity and population structure of the endemic Azorean juniper,Juniperus brevifolia (Seub.) Antoine,inferred from SSRs and ISSR markers

Juniperus brevifolia is an important woody species endemic of the Azores archipelago (Portugal), found from coastal to mountain environments. Due to colonization and grazing pressure this species has suffered fragmentation, leading to extinction in one island and being threatened in others. The genetic diversity and population structure of J. brevifolia populations was studied to provide guidelines for restoration and conservation programmes. Nuclear Single Sequence Repeats (nSSR) from Juniperus communis and Juniperus przewalskii and Inter Simple Sequence Repeats (ISSR) were tested and results compared to test the transferability of the microsatellites to J. brevifolia. Samples from ten populations over three islands, divided as coastal, mid-altitude and mountain were analysed. Both marker systems revealed results statistically and strongly correlated with each other, and not dependent on population sample size. We observed positive fixation indexes, moderate to high levels of genetic diversity (h = 0.415 for nSSR and h = 0.245 for ISSR), low to moderate φ_pt genetic differentiation (0.070 for nSSR and 0.129 for ISSR) and high gene flow (N_m > 2.432). Regarding Nei's genetic distance the coastal communities clustered together reflecting the phenotypic plasticity, but no specific clustering was observed regarding φ_pt values. Therefore no populations with substantial genetic differentiation were identified, once the diversity is mostly observed within populations. However it is advised the continuous monitoring of J. brevifolia.     

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.     

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     

New records for the shallow-water chiton fauna (Mollusca,Polyplacophora) of the Azores (NE Atlantic)

Published records, original data from recent field work on all of the islands of the Azores (NE Atlantic), and a revision of the entire mollusc collection deposited in the Department of Biology of the University of the Azores (DBUA) were used to compile a checklist of the shallow-water Polyplacophora of the Azores. Lepidochitona cf. canariensis and Tonicella rubra are reported for the first time for this archipelago, increasing the recorded Azorean fauna to seven species.     

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.     

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.     

Ranking protected areas in the Azores using standardised sampling of soil epigean arthropods

Nineteen areas in seven of the nine Azorean islands were evaluated for species diversity and rarity based on soil epigean arthropods. Fifteen out of the 19 study areas are managed as Natural Forest Reserves and the remaining four were included due to their importance as indigenous forest cover. Four of the 19 areas are not included in the European Conservation network, NATURA 2000. Two sampling replicates were run per study area, and a total of 191 species were collected; 43 of those species (23%) are endemic to the archipelago and 12 have yet to be described. To produce an unbiased multiple-criteria index (importance value for conservation, IV-C) incorporating diversity and rarity based indices, an iterative partial multiple regression analysis was performed. In addition, an irreplaceability index and the complementarity method (using both optimisation and heuristic methods) were used for priority-reserves analyses. It was concluded that at least one well-managed reserve per island is absolutely necessary to have a good fraction of the endemic arthropods preserved. We found that for presence/absence data the suboptimal complementarity algorithm provides solutions as good as the optimal algorithm. For abundance data, optimal solutions indicate that most reserves are needed if we want that at least 50% of endemic arthropod populations are represented in a minimum set of reserves. Consistently, two of the four areas not included in the NATURA 2000 framework were considered of high priority, indicating that vascular plants and bird species used to determine NATURA 2000 sites are not good surrogates of arthropod diversity in the Azores. The most irreplaceable reserves are those located in older islands, which indicates that geological history plays an important role in explaining faunal diversity of arthropods in the Azores. Based both on the uniqueness of species composition and high species richness, conservation efforts should be focused on the unmanaged Pico Alto region in the archipelago’s oldest island, Santa Maria.     

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.     

Parasite island syndromes in the context of nidicolous ectoparasites: Fleas (Insecta: Siphonaptera) in wild passerine birds from Azores Archipelago

Island syndrome, previously established for isolation process of insular vertebrates' populations, have been adapted to insular parasites communities, termed parasite island syndromes. In this work, were studied for the first time the insular syndromes for nidicolous ectoparasites of the bird species, Turdus merula, Sylvia atricapilla, Fringilla coelebs and Erithacus rubecula from Azores and the mainland Portugal. Flea species were only recorded on Azorean birds, namely Dasypsyllus gallinulae and Ctenocephalides felis felis, known as not host-specific parasites. In the absence of shared flea species between mainland and islands birds, a comparison among our fleas prevalence to Azores Islands and mainland fleas prevalence, recorded to others European studies, showed that Azorean host populations undergo higher prevalence than the mainland one. This result was consistent with parasite island syndromes predictions recorded to ectoparasites, hippoboscid flies and chewing lice, that fleas have higher prevalence on the Azores Islands compared to mainland Portugal. However, our results provide a new perspective to parasite island syndromes assumptions, namely in the context of nidicolous ectoparasites that spend only brief periods on the hosts' body.     

Island size, isolation, or interspecific competition? The breeding distribution of the Parus guild in the Danish archipelago

The Parus guild (Parus spp., Sitta, Certhia, and Regulus) is distributed as a complex mosaic within the Danish archipelago, with from one to eight species on different islands. We assessed the roles of island isolation, island size, and interspecific competition in determining the breeding species compositions of this guild on 53 Danish islands. Small, isolated islands supported fewer species than larger, nearshore islands. These effects, however, were largely restricted to a few sedentary species (P. cristatus, P. palustris, S. europaea) that are known to be poor dispersers/colonizers. In some cases, these three species were also absent from large, nearshore islands with suitable habitat, suggesting that habitat availability was not always responsible for the absence of a species. Monte Carlo simulations suggested that the pattern of species presence/absence was not a result of interspecific interactions. Thus, although a number of previous studies have documented interspecific competition among members of the Parus guild, our results suggest that such competition is not responsible for the unusual pattern of species distribution within the Danish archipelago. Received: 28 October 1996 / Accepted: 7 February 1997     

Parthenogenesis did not consistently evolve in insular populations of Ischnura hastata (Odonata,Coenagrionidae)

1. The evolutionary advantages that have driven the evolution of sex are still very much debated, and a number of benefits of parthenogenesis over sexual reproduction have been proposed. In particular, parthenogenetic individuals are thought to exhibit higher probabilities of establishment following arrival in new, isolated habitats such as islands. 2. One notable example of parthenogenesis occurring in islands is the damselfly Ischnura hastata, an American species that has colonised the Azores archipelago, where the populations consist only of females. This is the only known example of parthenogenesis within the insect order Odonata. 3. Here, two island populations of I. hastata were studied, one in the Galapagos and one in Cuba, to test whether island colonisation is consistently associated with parthenogenesis in this species. Field capture–mark–recapture studies and laboratory rearing of field‐collected eggs were undertaken in both areas. 4. Sex ratios in the field were found to be heavily female‐biased among mature individuals; however, fertility rates of field‐collected eggs were high, and the sex ratios in the laboratory did not differ from 1 : 1. Data from laboratory rearing showed that shorter larval development times and shorter adult life spans in males result in protandry, which might explain the skewed sex ratios in the field. 5. These findings are consistent with sex differences in key demographic parameters which could predispose I. hastata to parthenogenesis. However, the Azores population of I. hastata remains the only documented case of asexual reproduction in this insect group.     

Seasonal surveillance confirms the range expansion of Aedes japonicus japonicus (Theobald) (Diptera: Culicidae) to the Hawaiian Islands of Oahu and Kauai

The Asian bush mosquito, Aedes japonicus japonicus (Theobald) was not known to occur in the Hawaii archipelago until it was identified on the island of Hawaii in 2003. This mosquito species remained undetected on the neighboring islands for 8?years before it was discovered at the Honolulu International Airport on Oahu in 2012. By 2015, four Ae. j. japonicus mosquitoes were collected in the western mountains of Oahu and one was collected in the central mountains of Kauai. The collection of this invasive mosquito species across the neighboring Hawaiian Islands of Oahu and Kauai indicated the need for increased seasonal surveillance on these islands. Following nearly four years of surveillance, Ae. j. japonicus was also confirmed to occur in the eastern mountains of Oahu and in the central mountainous region of Kauai. To expand the knowledge of the spread of invasive mosquitoes species further surveillance is necessary to identify all possible areas where populations of Ae. j. japonicus and other invasive mosquito species occur in Hawaiian archipelago.     

Larval aquatic and terrestrial mites infesting parthenogenetic <Emphasis Type="Italic">Ischnura hastata</Emphasis> (Odonata: Coenagrionidae) from the Azores islands

We report here the prevalence of parasitism by water mites (Arrenurus sp.) and terrestrial mites (Leptus killingtoni) on parthenogenetic Ischnura hastata (Odonata: Coenagrionidae) from the Azores islands. Leptus killingtoni was only found on the island of Pico, and the prevalence of infestation was highly variable among the different ponds studied, ranging from 0 to 41%. Leptus killingtoni was observed on three of the four odonate species from the archipelago: I. hastata, I. pumilio, and Sympetrum fonscolombii, all of them new hosts for this species. Aquatic mites have been found parasitizing I. hastata females on the island of São Miguel. The prevalence of mite parasitism by Arrenurus sp. on I. hastata was very low, ranging from 12% (2003) to 1% (2008), and in most of the studied ponds, no mites were found attached to females. Although I. hastata coexists with a sexual congener species in the Azores (I. pumilio), they are syntopic in only a small fraction of ponds. Therefore, a comparison between I. hastata and I. pumilio was insufficient to test the predictions of the Red Queen Hypothesis, and further research on parasitism rates in both species needs to be done. In any case, the low prevalence of mite parasitism found in the Azores, coupled with the fact that most of the populations in the archipelago are almost free from competitors and predators, could explain the persistence of these I. hastata parthenogenetic populations, despite their low levels of genetic variation.     

Diversity of drosophilid flies on Kume-jima, a subtropical island: comparison with diversity on Iriomote-jima

The drosophilid fauna was studied on Kume‐jima, a subtropical island located in the central part of the Ryukyu archipelago, and compared with the fauna of Iriomote‐jima located near the south‐western end of the archipelago. The number of species collected from Kume‐jima was 37, much fewer than that recorded from Iriomote‐jima (95 species). The number of subtropical species was particularly reduced on Kume‐jima, possibly owing either to this island being more distantly located from the sources of subtropical species (e.g. Taiwan) than Iriomote‐jima and/or to winter temperature on Kume‐jima being a little lower (by approximately 1.5°C). The number of fungus‐feeders was also much reduced on Kume‐jima, but the number of fruit‐feeders was only slightly reduced. On Kume‐jima, fungi seem to be less abundant because forests are smaller, resulting in a smaller number of fungus‐feeders. Habitat selection and seasonality were analyzed for species collected using “retainer” type traps baited with banana. For species occurring on both islands, habitat selection differed little between the two islands, whereas the seasonality of some species differed markedly between the two islands.     

Spiders on a Hot Volcanic Roof: Colonisation Pathways and Phylogeography of the Canary Islands Endemic Trap-Door Spider Titanidiops canariensis (Araneae,Idiopidae)

Studies conducted on volcanic islands have greatly contributed to our current understanding of how organisms diversify. The Canary Islands archipelago, located northwest of the coast of northern Africa, harbours a large number of endemic taxa. Because of their low vagility, mygalomorph spiders are usually absent from oceanic islands. The spider Titanidiops canariensis, which inhabits the easternmost islands of the archipelago, constitutes an exception to this rule. Here, we use a multi-locus approach that combines three mitochondrial and four nuclear genes to investigate the origins and phylogeography of this remarkable trap-door spider. We provide a timeframe for the colonisation of the Canary Islands using two alternative approaches: concatenation and species tree inference in a Bayesian relaxed clock framework. Additionally, we investigate the existence of cryptic species on the islands by means of a Bayesian multi-locus species delimitation method. Our results indicate that T. canariensis colonised the Canary Islands once, most likely during the Miocene, although discrepancies between the timeframes from different approaches make the exact timing uncertain. A complex evolutionary history for the species in the archipelago is revealed, which involves two independent colonisations of Fuerteventura from the ancestral range of T. canariensis in northern Lanzarote and a possible back colonisation of southern Lanzarote. The data further corroborate a previously proposed volcanic refugium, highlighting the impact of the dynamic volcanic history of the island on the phylogeographic patterns of the endemic taxa. T. canariensis includes at least two different species, one inhabiting the Jandia peninsula and central Fuerteventura and one spanning from central Fuerteventura to Lanzarote. Our data suggest that the extant northern African Titanidiops lineages may have expanded to the region after the islands were colonised and, hence, are not the source of colonisation. In addition, T. maroccanus may harbour several cryptic species.     

Distribution and taxonomy of shallow marine Ostracods from Northern Socotra Island (Indian Ocean) - Yemen

Forty-seven species belonging to 30 genera of ostracods have been identified from Recent shallow marine sediments collected along the northern coastline of the island of Socotra (Indian Ocean). The systematics and distribution of these ostracods are discussed. Three genera, Loxoconcha, Bosasella, and Xestoleberis are found to have high species diversity. The species Loxoconcha ghardaqensis, Moosella striata are best documented in the Red Sea and Gulf of Aden while Kotoracythere inconspicua and Triebelina sertata were found to be the most widespread species. The fauna shows a close affinity to other ostracod assemblages of the Indo-Pacific region, East African coast and Red Sea.