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.     

New species of Rissoidae (Mollusca,Gastropoda) from the Archipelago of the Azores (northeast Atlantic) with an updated regional checklist for the family

Four new species of shallow-water marine gastropods belonging to the family Rissoidae are described from the Archipelago of the Azores: Setia alexandrae sp. n., Setia ermelindoi sp. n., Setia netoae sp. n., and Manzonia martinsi sp. n. These novelties increase the regional rissoid fauna to 39 species, of which 29 live in shallow-water habitats. A list of the species of Rissoidae from the Azores is presented based on data from the literature and new material examined.     

Population genetic structure and conservation of the Azorean tree Prunus azorica (Rosaceae)

Prunus azorica is an endangered tree endemic to the Azores Archipelago, considered as a top priority species for conservation. Although propagation measures have been studied in detail, and a broad phylogeographic study on P. lusitanica was recently published, a detailed population genetics study devoted to Azorean taxon was lacking. To determine extant patterns of population genetic structure in P. azorica, we analysed eight populations from the five Azorean islands where the species presently occurs and the only extant individual from Flores Island. We also included samples of P. lusitanica subsp. hixa from the Canary Islands and Madeira, and of P. lusitanica subsp. lusitanica from mainland Portugal. Genotyping was undertaken for eight nuclear microsatellite polymorphic loci specifically isolated for P. azorica. Accessions of the different geographic regions were used to sequence ITS and trnL DNA regions. Regarding SSRs, the number of alleles ranged from 5 to 37 (mean = 12.6) per locus and from 2 to 64 per population (mean = 24). Our analysis showed a clear separation between samples from the Azores and those from other regions. Overall, São Miguel populations seemed to encompass the majority of the variability found within the archipelago. Regarding the Azorean populations only, the highest percentage of genetic variation was found within populations (92 %). Still, about 7 % of the variation was found among populations within islands. Expected heterozygosity ranged from values near 0 in the most depauperate populations up to 0.18. With a few exceptions, the level of differentiation between Azorean populations was generally low and gene flow was clearly above 1. Analysis of ITS sequences also detected differences between the Azores and the remaining regions but the trnL region did not reveal any variation. The genetic identity of P. azorica was recognised and thus should be preserved; however, the present results suggest that the Azorean taxon should be reinstated at the subspecies level.     

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.     

The genus Huperzia (Lycopodiaceae) in the Azores and Madeira

The taxonomy and nomenclature of the genus Huperzia Bernh. in the Azores and Madeira have been reviewed. Plants collected in the Azores and Madeira were characterized morphologically. The independence between two endemic species common to Madeira and the Azores Islands –Huperzia suberecta (Lowe) Tardieu and Huperzia dentata (Herter) Holub – is clearly shown. A clear‐cut morphological separation between these taxa and Huperzia selago (L.) Bernh. ex Schrank & Mart. of continental Europe is established. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 158 , 522–533.     

Preliminary observations on the benthic marine algae of the Gorringe seabank (northeast Atlantic Ocean)

Examination of marine samples collected in 2006 from the Gettysburg and Ormonde seamounts on the Gorringe seabank southwest of Portugal has revealed 29 benthic Chlorophyta, Phaeophyceae (Ochrophyta), and Rhodophyta that were identified provisionally to genus and to species. Combining lists for the present and a previous expedition brings the total of algae thus far recorded to 48. The brown alga Zonaria tournefourtii and the red alga Cryptopleura ramosa were the most abundant species in the present collections. The kelp Laminaria ochroleuca was present only in the Gettysburg samples while Saccorhiza polyschides was observed only on the Ormonde seamount. Comparisons with the benthic marine algae recorded on seamounts in the mid-Atlantic Azores archipelago show features in common, notably kelp forests of L. ochroleuca at depths below 30 m and Z. tournefortii dominance in shallower waters.     

Visual observations of fish from seamounts of the Southern Azores region (the Atlantic Ocean)

Results of observations from the underwater inhabited device “Sever-2” of the behavior and distribution of fish on seamounts of the Azores region up to a depth of 1200 m are provided. In the natural habitat, 52 species of fish of 46 genera belonging to 36 families were recorded. In the ichthyofauna of the studied region, bottom (37.0%) and near-bottom (29.2%) species dominate; the proportion of off-bottom and off-bottom-pelagic species is 18.0 and 12.4%, respectively, and pelagic species account for only 3.4%. Pelagic species Scomber japonicus and Trachurus picturatus; off-bottom-pelagic Beryx splendens, Lepidopus caudatus, and Aphanopus carbo; and near-bottom species Zenopsis conchifer, Macrorhamphosus scolapax, Antigonia capros, Capros aper, Anthias anthias, and Callanthias ruber dominate by numbers and the amount of biomass.     

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).     

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.     

Actiniaria and Ceriantharia of the Azores (Cnidaria Anthozoa)

The common shallow water species of sea anemones (Actiniaria) and tube anemones (Ceriantharia) of the Azores are listed. Eight species of sea anemones are mentioned, the species Cereus pedunculatus and Sagartia affinis being new records for the archipelago. Both species of Ceriantharia, namely Arachnanthus nocturnus and Pachycerianthus solitarius, are recorded from the Azores for the first time. Arachnanthus nocturnus is also recorded from the Cape Verde Islands and from Madeira for the first time. Communicated by H.-D. Franke     

Genetic and morphometric diversity of the goldcrest (Regulus regulus) populations in the Azores

We studied the genetic diversity and phylogeography of the goldcrest Regulus regulus from the archipelago of the Azores (North Atlantic Ocean) based on sequences of two mitochondrial genes (cytochrome b and the NADH dehydrogenase subunit 2) and one nuclear gene in the Z-chromosome (intron 9 of the aconitase 1) from 69 individuals, and 41 birds from the Canary Islands and continental Europe for outgroup comparison. To understand the level of concordance between the genetic data and possible morphometric variability, 197 adult living birds from the seven Azorean islands where the species breeds were analysed in terms of eight morphometric characters. Our results are in accordance with previous studies, indicating a recent expansion of goldcrests throughout the archipelago and a low divergence in relation to continental Europe. Within the Azores, there is evidence of historical and/or recent gene flow among the island's populations, revealing a lack of current genetic structure within the archipelago. Only goldcrests from Flores Island seem to be genetically distinct and showed significantly larger body mass and tarsus length than birds on the other islands.     

Parasitoids from Azores (Hymenoptera: Encyrtidae,Pteromalidae, Braconidae): potential use in integrated pest management against Ceratitis capitata (Diptera: Tephritidae)

Two hymenopterous parasitoids, Halticoptera patellana and Toxeumorpha nigricola, were recovered from Mediterranean fruit fly pupae on São Miguel Island, Azores. Both are new records for the Azores, and as Medfly parasitoids. Field-collected Tachinaephagus zealandicus was maintained on Medfly for 10 generations, but if it has potential as a biocontrol agent remains an open question.     

Ichthyoplankton of the Azores seamounts

Species composition and quantitative distribution of ichthyoplankton in the region of the oceanic seamounts of the Azores complex from 2000 to 2004 are considered. The spawning biomass of the main fishery item of this region—Atlantic jack Trachurus picturatus—is calculated. The ichthyoplankton of the Azores seamounts in the period under consideration included eggs and larvae of 60 fish species from 26 families. In the summer, 20 species of eggs and larvae from 11 families and, in winter, 57 species from 24 families were recorded. Larval Myctophidae and Gonastomatidae have the most mass in the plankton in both seasons. The eggs and larvae of commercial fish species (T. picturatus and Pacific mackerel Scomber japonicus) were recorded mainly in the cold period. The species and quantitative composition of ichthyoplankton of the South Azores seamounts is far more numerous than in the North Azores seamounts. The region of the seamounts of the South Azores complex, judging from the estimation of biomass of commercial fish with different methods, can be regarded as a reserve for the fishery of Russian vessels in the central eastern Atlantic.     

First record of the pencil cardinal Epigonus denticulatus (Perciformes: Epigonidae) in the Azores archipelago

The pencil cardinal Epigonus denticulatus is a small deep-water fish inhabiting continental slopes usually between 300 and 600 m depth. We report the first record of E. denticulatus in the Azores archipelago, where one specimen was found floating by fisherman off Faial island. Meristic and morphometric characters are in accordance with those reported for the species and molecular analyses further supported species identity. The record of E. denticulatus as a native species in the Azores increases the number of Epigonus species in the region to a total of three.     

Hybridization and speciation in the genusDryopteris (Pteridophyta:Dryopteridaceae) on Pico Island in the Azores

The cytology of wild hybrids ofDryopteris from Pico, Azores has been examined. This has confirmed the parentage of the hybrids, and provided evidence to support the view that the tetraploid speciesD. crispifolia has evolved in the Azores from two diploid species,D. aemula andD. azorica, by hybridization and polyploidy.     

Resolving the Azorean knot: a response to Carine & Schaefer (2010)

Carine & Schaefer (Journal of Biogeography, 2010, 37 , 77–89) suggest that the lack of past climate oscillations in the Azores may have contributed to the low plant endemism in this archipelago compared to that of the Canary Islands, a pattern they term the Azorean diversity enigma. Here we challenge their hypothesis, and discuss how the particular characteristics of the Azores may have driven current diversification patterns in this archipelago. We argue that the restricted number of Azorean endemic species and their wide distribution is explicable by the geological, geographical and ecological attributes of the archipelago. That is, the Azores are too young, too small, and too environmentally homogeneous to have hosted many in situ diversification events, so they do not host as many endemic species as other Macaronesian archipelagos, such as Madeira and especially the Canary Islands.     

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.     

Geography,geology and ecology influence population genetic diversity and structure in the endangered endemic Azorean <Emphasis Type="Italic">Ammi</Emphasis> (Apiaceae)

Three Azorean endemic Ammi species were initially described: Ammi trifoliatum (Wats.) Trel., Ammi seubertianum (Wats.) Trel. and Ammi huntii (Wats.) Trel. Many taxonomic changes have been conducted, and one to three species have been considered. Two species are currently accepted: A. trifoliatum, which occurs in almost every island, and A. seubertianum, with a narrower distribution. In this research, the population genetic diversity and structure of the Azorean Ammi species were assessed using eight specifically designed SSR markers. A wide sampling of A. seubertianum and A. trifoliatum was conducted in seven Azorean islands, and four A. huntii herbarium samples were also included to further contribute to the taxonomy of this genus in Azores. Flores populations showed the highest genetic diversity, while North of Topo, in São Jorge, showed the lowest. None of the populations analysed displayed signs of putative inbreeding. The population genetic structure analyses conducted partially provided support for the two currently accepted species, but other possible cryptic taxa may also be present. The complex clustering obtained seems to result from a combined action of geography, geology and ecology, and although island-specific genetic patterns were found, environmental conditions connected to different altitudes and the existence of micro-niches may also play an important role. A thorough morphological revision and ecophysiological studies should be conducted to clarify the number of endemic taxonomic entities present in the Azores archipelago.     

Physiological responses of Pterocladia and Gelidium (Gelidiales,Rhodophyta) from the Azores,Portugal

Manometric studies were conducted on Pterocladia capillacea, Gelidium latifolium and Gelidium spinulosum from the Azores, Portugal to determine optimal values of temperature, light and salinity for growth. Physiological responses were considered in relation to vertical distribution patterns of these species commonly observed throughout the Azores. Optimal parameters for the growth of Pterocladia capillacea, Gelidium latifolium and G. spinulosum were 17 to 25 °C, a photon flux density between 200 and 300 µmol m^–2 s^–1 and salinities of 25 to 35.     

Dendrochronological potential of the Azorean endemic gymnosperm Juniperus brevifolia (Seub.) Antoine

Tree-ring inter-annual pattern variation is crucial in dendrochronology, allowing the identification of possible limiting factors on growth. Thus, trees exposed to subtropical or tropical climates without a marked seasonality may show a low degree of interannual variation, impeding a straightforward dendroclimatological approach. Meanwhile, subtropical regions, and areas in transitional climates such as the Azores archipelago, are widely unexplored in terms of dendroclimatology, providing opportunities to work with endemic trees, including the dominant Azorean tree Juniperus brevifolia (Seub.) Antoine. To evaluate the dendrochronological potential of J. brevifolia, we analyzed tree-ring patterns, crossdating capabilities, and correlation with climate parameters. We sampled 48 individual trees from two natural populations (São Miguel and Terceira islands) using an increment borer. Besides, a Trephor tool was used to obtain wood microcores for micro-anatomical analysis. Although the transition between early and latewood was evident, partially indistinct ring boundaries and wedging rings were present in some cases, affecting the crossdating process, but not impeding the establishment of reliable ring-width chronologies. Following detrending, master chronologies were built and correlated with monthly temperature and precipitation data using the treeclim R package. The climate-growth relationships indicated negative correlations with late summer temperature in both populations. Considering our results and the importance of J. brevifolia as a dominant tree in the Azores natural forests, we conclude that it shows an acceptable potential for dendrochronological research. Thus, this study provides baseline information to help fill the knowledge gap regarding the climate-growth relationship of Azorean trees.