Ecological traits influence the current distribution and range of an island endemic bird

Aim  The aim of this paper is to investigate the causes of the current restricted distribution of a narrow-range endemic bird species, the Canary Islands stonechat, Saxicola dacotiae .
Location  Eastern islands of the Canary Islands archipelago.
Methods  We compared climatic patterns (temperature and rainfall), habitat and microhabitat structure, food availability during a full annual cycle, and the abundance of native avian competitors and predators inside and outside the species' range. Three study areas, located in similar habitats on nearby islands, were studied: northern Fuerteventura, close to the northern border of the species' range; southern Lanzarote, 22 km from the nearest site occupied by stonechats; and the Lobos islet, 10 km from the nearest occupied site and 2 km from the coast of Fuerteventura.
Results  The cover of suitable habitats (slopes with high cover of large shrubs, stony fields and ravines) and microhabitats (shrubs and boulders) and the abundance of arthropods during the breeding period of Canary Islands stonechats were lower outside than inside the species' range. Temperature, rainfall and the abundance of competitors and predators inside and outside the species' range did not differ significantly.
Main conclusions  Ecological requirements explaining the distribution of the Canary Islands stonechat within its range seem to be the main factor hindering its settlement on nearby islands. Geological and palaeoclimatic processes, as well as past and current human impact, could also have constrained the distribution of this narrow-range endemic bird species.     

Xenopsylla spp. (Siphonaptera: Pulicidae) in murid rodents from the Canary Islands: An update

The geographical and host distributions of Xenopsylla fleas parasitizing murid rodents on the Canary Islands have been reported. Three Xenopsylla species, X. cheopis, X. brasiliensis and X. guancha, have been detected on two rodents species, Mus musculus and Rattus rattus. X. guancha has been the most prevalent species detected, specifically on M. musculus, the most abundant rodent, but it has been detected only on three eastern islands, where the species is endemic. X. cheopis has been shown to be the most widely distributed species throughout the archipelago and the species most frequently found on R. rattus. X. brasiliensis has been shown to be the least prevalent Xenopsylla species, with the lowest geographical distribution on the Canary Islands and focused only on R. rattus. The detection of both X. cheopis and X. brasiliensis on the island of Lanzarote, and of X. guancha on the island of Fuerteventura and the islet of La Graciosa represents the first report of these species on those particular Canary Islands.     

Effects of climate change on the distribution of plant species and plant functional strategies on the Canary Islands

Aim

Oceanic islands possess unique floras with high proportions of endemic species. Island floras are expected to be severely affected by changing climatic conditions as species on islands have limited distribution ranges and small population sizes and face the constraints of insularity to track their climatic niches. We aimed to assess how ongoing climate change affects the range sizes of oceanic island plants, identifying species of particular conservation concern.

Location

Canary Islands, Spain.

Methods

We combined species occurrence data from single-island endemic, archipelago endemic and nonendemic native plant species of the Canary Islands with data on current and future climatic conditions. Bayesian Additive Regression Trees were used to assess the effect of climate change on species distributions; 71% (n = 502 species) of the native Canary Island species had models deemed good enough. To further assess how climate change affects plant functional strategies, we collected data on woodiness and succulence.

Results

Single-island endemic species were projected to lose a greater proportion of their climatically suitable area (x ̃ = −0.36) than archipelago endemics (x ̃ = −0.28) or nonendemic native species (x ̃ = −0.26), especially on Lanzarote and Fuerteventura, which are expected to experience less annual precipitation in the future. Moreover, herbaceous single-island endemics were projected to gain less and lose more climatically suitable area than insular woody single-island endemics. By contrast, we found that succulent single-island endemics and nonendemic natives gain more and lose less climatically suitable area.

Main Conclusions

While all native species are of conservation importance, we emphasise single-island endemic species not characterised by functional strategies associated with water use efficiency. Our results are particularly critical for other oceanic island floras that are not constituted by such a vast diversity of insular woody species as the Canary Islands.     

Evolution of an invasive rodent on an archipelago as revealed by molar shape analysis: the house mouse in the Canary Islands

Aim The aim of this paper is to identify the patterns in the morphological differentiation in Canary Island mice, based on fossil and modern samples. In order to achieve this, the mouse species present on the archipelago were first compared with a set of continental mice. The differences between the continental and Canary Island samples, and among the Canary Island samples, provide insights into the processes of colonization and the subsequent insular evolution. Location Canary archipelago. Methods An outline analysis based on Fourier transformation was used to quantify shape differences between lower molars. Together with the fossil and modern Canary Island samples, a reference set of genotyped continental populations of the commensal Mus musculus and the wild Mus spretus was used for comparison. Results The morphometric analysis showed that all the mouse specimens from the Canary Islands and Cape Verde belonged to Mus musculus domesticus. Lower molars of extant mice from La Gomera, El Hierro, Gran Canaria, Tenerife, and to a lesser degree from Lanzarote, were similar to those of genotyped M. m. domesticus from the continent, while teeth of extant mice from Fuerteventura were more divergent. Fossil mice from Fuerteventura were very similar to the extant representatives on this island, and similar to the fossil mice on the nearby islands of Lobos and La Graciosa. Main conclusions The mouse present on the Canary archipelago has been identified as the house mouse M. m. domesticus. Based on the shape of the lower molar, the Canary Island mice are divergent from the continental ones, but the degree of divergence varies with the geography of the archipelago. Overall, populations from eastern islands are more divergent from the continental mice than populations from western ones. Fossil populations indicate that this situation was established several centuries ago. Two main factors may have contributed to this pattern: the appearance of different types of environment on the islands since the successful settlement of the mouse, and/or the number of subsequent introductions of continental individuals via shipping.     

The phylogeography of the darkling beetle,Hegeter politus,in the eastern Canary Islands.

We investigated the phylogeography of Hegeter politus, a saprophagous, flightless darkling beetle endemic to the eastern Canary Islands, using a fragment of the mitochondrial COI gene. Distance and parsimony based gene trees of the mitotypes identified revealed a striking association between mitotype clades and sampling locations. The branching order of the clades suggested that the colonization of the islands by Hegeter politus proceeded from the southern part of Fuerteventura in a north-northeast direction to Lanzarote and the smaller islands. Based on this, a colonization scenario compatible with the reported geological ages and volcanisms of the various parts of the islands has been proposed. The high divergence of the beetles collected from the extreme south of Fuerteventura (the Jandía peninsula) from all other samples has led us to propose that they may be from a new species that has not been described previously. The ecological isolation of Jandía from the rest of Fuerteventura by the sand dunes that cover its narrow isthmus in the north, and the existence of many plant and animal endemisms unique to Jandía, lend supportive evidence to our proposal. The similarities between the evolution of island endemics in the Hawaiian and Canary archipelagos have been discussed. We conclude that many endemics in the Canary archipelago, like the Hawaiian Islands, are most likely to have originated from post-colonization differentiation and divergence.     

Complex population genetic structure in the endemic Canary Island pine revealed using chloroplast microsatellite markers

The Canary archipelago, located on the northwestern Atlantic coast of Africa, is comprised of seven islands aligned from east to west, plus seven minor islets. All the islands were formed by volcanic eruptions and their geological history is well documented providing a historical framework to study colonization events. The Canary Island pine (Pinus canariensis C. Sm.), nowadays restricted to the westernmost Canary Islands (Gran Canaria, Tenerife, La Gomera, La Palma and El Hierro), is considered an old (Lower Cretaceous) relic from an ancient Mediterranean evolutionary centre. Twenty seven chloroplast haplotypes were found in Canary Island pine but only one of them was common to all populations. The distribution of haplotypic variation in P. canariensis suggested the colonization of western Canary Islands from a single continental source located close to the Mediterranean Basin. Present-day populations of Canary Island pine retain levels of genetic diversity equivalent to those found in Mediterranean continental pine species, Pinus pinaster and Pinus halepensis. A hierarchical analysis of variance (AMOVA) showed high differentiation among populations within islands (approximately 19%) but no differentiation among islands. Simple differentiation models such as isolation by distance or stepping-stone colonization from older to younger islands were rejected based on product-moment correlations between pairwise genetic distances and both geographic distances and population-age divergences. However, the distribution of cpSSR diversity within the islands of Tenerife and Gran Canaria pointed towards the importance of the role played by regional Pliocene and Quaternary volcanic activity and long-distance gene flow in shaping the population genetic structure of the Canary Island pine. Therefore, conservation strategies at the population level are strongly recommended for this species.Communicated by D.B. NealeA. Gómez and S.C. González-Martínez as joint authors     

Predicting potential distributions of invasive species: the exotic Barbary ground squirrel in the Canarian archipelago and the west Mediterranean region

This contribution aimed to predict the invasive Barbary ground squirrel (Atlantoxerus getulus) potentiality for invading the Canary Islands and western Mediterranean region, by determining firstly the climatic suitable areas in its native range and secondly, using presence data in the invaded range. Nineteen environmental variables submitted to a Principal Components Analysis selected those variables with higher factor loadings, which represent the main environmental conditions of the Northern African region (temperature in the coldest quarter, seasonal temperature, precipitation in the coldest quarter, temperature in the wettest quarter). After selecting hundred times more pseudo-absence points than presence observations (n = 6600 at a 0.083° resolution), Generalized Additive Models and Single-hidden-layer Neural Networks fitted in R were used to calibrate the model. Model results were extrapolated for the Canary Islands and the western Mediterranean region. In order to select between the two techniques, we calculated three accuracy measures (specificity, sensitivity and AUC) after using a Jack-knifing procedure and models were repeated ten times. The GAM model was less accurate than the NN model. Suitable areas did not have mean temperatures in the coldest quarter lower than −5°C and precipitation in the coldest quarter higher than 300 mm, respectively. We predicted favorable climatic areas across almost all the Maghreb, the European western Mediterranean region and in all the Canary Islands. Nevertheless, the seven islands differed significantly in the mean favorability scores, with El Hierro, Lanzarote and Gran Canaria being the most suitable. Same methodological analysis was applied to predict A. getulus distribution in other Canarian islands based on presence data from the invaded Fuerteventura. In this case, only Lanzarote and Gran Canaria appeared to be climatically suitable for the species. Our predictive model is an applicable tool to establish the invasive potential of A. getulus and to prioritize management strategies, within and outside the Canarian archipelago, to impede the expansion of this invasive squirrel out of Fuerteventura Island.     

David Fairchild expeditions to the Canary Islands: Plant collections and research outcomes

Dr. David Fairchild (1869–1954) made four field trips to the Canary Islands. They took place in 1903 (supported by Barbour Lathrop) and in 1925, 1926, and 1927 (supported by Allison V. Armour). They represented the first extensive plant exploration expeditions ever undertaken by United States botanists targeting this archipelago. At least two of the trips were joined by other biologists from several disciplines. Four of the seven Canary Islands (Gran Canaria, Lanzarote, La Palma, and Tenerife) were visited. The four expeditions yielded eight publications, over 315 photographs, and 89 germplasm accessions (73 species). Two of these publications focused on invertebrates, one on plant mosaic viruses, and two on the pollination biology and propagation of Ficus auriculata (Moraceae). The remaining three works provided accounts on the natural history and plants of the islands and on the cropping systems and the rural life of the archipelago.     

Molecular evidence for adaptive radiation of Micromeria Benth. (Lamiaceae) on the Canary Islands as inferred from chloroplast and nuclear DNA sequences and ISSR fingerprint data

The Canary Islands have been a focus for phylogeographic studies on the colonization and diversification of endemic angiosperm taxa. Based on phylogeographic patterns, both inter island colonization and adaptive radiation seem to be the driving forces for speciation in most taxa. Here, we investigated the diversification of Micromeria on the Canary Islands and Madeira at the inter- and infraspecific level using inter simple sequence repeat PCR (ISSR), the trnK-Intron and the trnT-trnL-spacer of the cpDNA and a low copy nuclear gene. The genus Micromeria (Lamiaceae, Mentheae) includes 16 species and 13 subspecies in Macaronesia. Most taxa are restricted endemics, or grow in similar ecological conditions on two islands. An exception is M. varia, a widespread species inhabits the lowland scrub on each island of the archipelago and could represent an ancestral taxon from which radiation started on the different islands. Our analyses support a split between the "eastern" islands Fuerteventura, Lanzarote and Gran Canaria and the "western" islands Tenerife, La Palma and El Hierro. The colonization of Madeira started from the western Islands, probably from Tenerife as indicated by the sequence data. We identified two lineages of Micromeria on Gomera but all other islands appear to be colonized by a single lineage, supporting adaptive radiation as the major evolutionary force for the diversification of Micromeria. We also discuss the possible role of gene flow between lineages of different Micromeria species on one island after multiple colonizations.     

Die Vogelwelt von La Graciosa

The terrestrial birds of La Graciosa The small island La Graciosa is part of an archipelago north of Lanzarote, the northern‐ and easternmost of the Canary Islands. Its habitats are mainly semi‐desert made up of sand dunes covered with dry shrubbery. They house only a small number of terrestrial birds, among them endemic species and subspecies of the Canary Islands. Birds are more common around the settlement of Caleta del Sebo where fresh water and more plant species are available.     

Palaeo‐islands as refugia and sources of genetic diversity within volcanic archipelagos: the case of the widespread endemic Canarina canariensis (Campanulaceae)

Geographical isolation by oceanic barriers and climatic stability has been postulated as some of the main factors driving diversification within volcanic archipelagos. However, few studies have focused on the effect that catastrophic volcanic events have had on patterns of within‐island differentiation in geological time. This study employed data from the chloroplast (cpDNA haplotypes) and the nuclear (AFLPs) genomes to examine the patterns of genetic variation in Canarina canariensis, an iconic plant species associated with the endemic laurel forest of the Canary Islands. We found a strong geographical population structure, with a first divergence around 0.8 Ma that has Tenerife as its central axis and divides Canarian populations into eastern and western clades. Genetic diversity was greatest in the geologically stable ‘palaeo‐islands’ of Anaga, Teno and Roque del Conde; these areas were also inferred as the ancestral location of migrant alleles towards other disturbed areas within Tenerife or the nearby islands using a Bayesian approach to phylogeographical clustering. Oceanic barriers, in contrast, appear to have played a lesser role in structuring genetic variation, with intra‐island levels of genetic diversity larger than those between‐islands. We argue that volcanic eruptions and landslides after the merging of the palaeo‐islands 3.5 Ma played key roles in generating genetic boundaries within Tenerife, with the palaeo‐islands acting as refugia against extinction, and as cradles and sources of genetic diversity to other areas within the archipelago.     

A new marine interstitial ingolfiellid (Crustacea,Amphipoda, Ingolfiellidea) from Tenerife and Hierro

Ingolfiella canariensis n. sp., from coarse sand and gravel in the mediolittoral zone of Tenerife and Hierro, Canary Islands, is described. The new species shares supposedly apomorphous characters with species from comparable habitats from the Andaman Islands, Bermuda and Curaçao (Netherlands Antilles). The female of Ingolfiella similis Rondé-Broekhuizen & Stock, 1987, from Fuerteventura is also described.     

Ilex Canariensis Poir. (Aquifoliaceae) Post-dispersal Seed Predation in the Canary Islands

Alien species have many negative effects on insular ecosystems worldwide. We investigated Ilex canariensis post-dispersal seed predation by introduced rats (Rattus spp.) in relict forests of the Canary Islands at different spatial scales: among microhabitats within the same forest, among forest types within the same island, and among different islands of the archipelago. Seed predation intensity was very high (>70%) in all cases considered, irrespective of the spatial scale. We did not find significant differences between forest interior, edges or gaps, as well as between different forest types in four islands of the archipelago. Comparatively low predation intensity was found in El Hierro island, where more than 50% of the seeds survived at the end of the experiment, while highest seed predation was observed in Tenerife island. It is concluded that post-dispersal seed predation by rats, due to its extent and intensity, could have an important effect on Ilex canariensis recruitment, especially in successional areas where this light tolerant tree can naturally establish.     

On the occurrence of egg masses of the diamond-shaped squid Thysanoteuthis rhombus Troschel, 1857 in the subtropical eastern Atlantic (Canary Islands). A?potential commercial species?

Data on opportunistic sightings of diamond-shaped squid Thysanoteuthis rhombus egg masses in the Canary Islands (Atlantic Ocean) are presented. A total of 16 egg masses of this species were recorded and photographed from 2000 to 2010 around the western islands of the archipelago (El Hierro, Tenerife and La Gomera). These data reveal the existence of an important spawning area for diamond-shaped squid around the Canary Islands, in subtropical east Atlantic waters. We provide preliminary data for the potential development of an artisanal fishery focused on this species, and a discussion on its potential impacts on the marine ecosystem.     

Radiating on oceanic islands: patterns and processes of speciation in the land snail genus Theba (Risso 1826)

Island radiations have played a major role in shaping our current understanding of allopatric, sympatric and parapatric speciation. However, the fact that species divergence correlates with island size emphasizes the importance of geographic isolation (allopatry) in speciation. Based on molecular and morphological data, we investigated the diversification of the land snail genus Theba on the two Canary Islands of Lanzarote and Fuerteventura. Due to the geological history of both islands, this study system provides ideal conditions to investigate the interplay of biogeography, dispersal ability and differentiation in generating species diversity. Our analyses demonstrated extensive cryptic diversification of Theba on these islands, probably driven mainly by non-adaptive allopatric differentiation and secondary gene flow. In a few cases, we observed a complete absence of gene flow among sympatrically distributed forms suggesting an advanced stage of speciation. On the Jandía peninsula genome scans suggested genotype-environment associations and potentially adaptive diversification of two closely related Theba species to different ecological environments. We found support for the idea that genetic differentiation was enhanced by divergent selection in different environments. The diversification of Theba on both islands is therefore best explained by a mixture of non-adaptive and adaptive speciation, promoted by ecological and geomorphological factors.     

The evolutionary history of Afrocanarian blue tits inferred from genomewide SNPs

A common challenge in phylogenetic reconstruction is to find enough suitable genomic markers to reliably trace splitting events with short internodes. Here, we present phylogenetic analyses based on genomewide single‐nucleotide polymorphisms (SNPs) of an enigmatic avian radiation, the subspecies complex of Afrocanarian blue tits (Cyanistes teneriffae). The two sister species, the Eurasian blue tit (Cyanistes caeruleus) and the azure tit (Cyanistes cyanus), constituted the out‐group. We generated a large data set of SNPs for analysis of population structure and phylogeny. We also adapted our protocol to utilize degraded DNA from old museum skins from Libya. We found strong population structuring that largely confirmed subspecies monophyly and constructed a coalescent‐based phylogeny with full support at all major nodes. The results are consistent with a recent hypothesis that La Palma and Libya are relic populations of an ancient Afrocanarian blue tit, although a small data set for Libya could not resolve its position relative to La Palma. The birds on the eastern islands of Fuerteventura and Lanzarote are similar to those in Morocco. Together they constitute the sister group to the clade containing the other Canary Islands (except La Palma), in which El Hierro is sister to the three central islands. Hence, extant Canary Islands populations seem to originate from multiple independent colonization events. We also found population divergences in a key reproductive trait, viz. sperm length, which may constitute reproductive barriers between certain populations. We recommend a taxonomic revision of this polytypic species, where several subspecies should qualify for species rank.     

Evolution and biogeography of the genus Tarentola (Sauria: Gekkonidae) in the Canary Islands,inferred from mitochondrial DNA sequences

Sequences from fragments of the 12S ribosomal RNA and cytochrome b mitochondrial genes were used to analyze phylogenetic relationships among geckos of genus Tarentola from the Canary Islands. A surprisingly high level of within island differentiation was found in T. delalandii in Tenerife and T. boettgeri in Gran Canaria. Molecular differentiation between populations of T. angustimentalis on Lanzarote and Fuerteventura, and between Moroccan and Iberian Peninsula T. mauritanica, also indicate that at least two subspecies should be recognized within each of them. Phylogenetic relationships among these species reveals a higher level of differentiation and a more complex colonization pattern than those found for the endemic genus Gallotia. Lack of evidence for the presence of T. boettgeri bischoffi on the island of Madeira does not seem to support the origin of T. delalandii, T. gomerensis and the canarian subspecies of T. boettgeri from this island, whereas molecular data confirms that T. angustimentalis is a sister species of the continental T. mauritanica. Several independent colonization events from the continent and the extinction of some species are probably responsible for the current distribution of Tarentola in the Canary Islands.     

Explanations for bird species range size: ecological correlates and phylogenetic effects in the Canary Islands

Aim To explore the determinants of island occupancy of 48 terrestrial bird species in an oceanic archipelago, accounting for ecological components while controlling for phylogenetic effects. Location The seven main islands of the Canary archipelago. Methods We obtained field data on population density, habitat breadth and landscape distribution in Tenerife, Fuerteventura and La Palma, aiming to sample all available habitats and the gradient of altitudes. In total, 1715 line transects of 0.5 km were carried out during the breeding season. We also reviewed the literature for data on occupancy, the distance between the Canary Islands and the nearest distribution border on the mainland, body size and endemicity of the 48 terrestrial bird species studied. Phylogenetic eigenvector regression was used to quantify (and to control for) the amount of phylogenetic signal. Results The two measurements of occupancy (number of occupied islands or 10 × 10 km UTM squares) were tightly correlated and produced very similar results. The occupancy of the terrestrial birds of the Canary Islands during the breeding season had a very low phylogenetic effect. Species with broader habitat breadth, stronger preferences for urban environments, smaller body size, and a lower degree of endemicity had a broader geographical distribution in the archipelago, occupying a larger number of islands and 10 × 10 UTM squares. Main conclusions The habitat‐generalist species with a tolerance for novel urban environments tend to be present on more islands and to occupy a greater area, whereas large‐sized species that are genetically differentiated within the islands are less widespread. Therefore, some properties of the ranges of these species are explicable from basic biological features. A positive relationship of range size with local abundance, previously shown in continental studies, was not found, probably because it relies on free dispersal on continuous landmasses, which may not be applicable on oceanic islands.     

Evolution and island endemism of morphologically cryptic Baetis and Cloeon species (Ephemeroptera,Baetidae) on the Canary Islands and Madeira

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Diversity, abundance and depth distribution of demersal deep-water fishes off Lanzarote and Fuerteventura, Canary Islands

A total of 168 bottom longline operations were carried out at depths between 72 and 1102 m during four cruises off Lanzarote and Fuerteventura, Canary Islands. One hundred and twenty-eight operations were successful, resulting in the capture of 1167 fishes belonging to 55 species. Clear variations in fish total weight and number, species richness and diversity, and the relative abundance of single species occurred among five different depth zones between 100 and 1100 m. The particular platform topography of the Canary Islands may affect the density and diversity of slope-dwelling fishes. Whereas fish body length and weight showed an overall increase with depth, no such trends were found at the level of individual species. One species, Lepidopus caudalus , even revealed a bigger-shallower trend. Among three species of trichiurids, depth distribution patterns were detected that may reflect vertical space partitioning. In October 1995, an unusually high number of Mora moro with advanced gonadal maturity were collected off eastern Fuerteventura at 997 m depth. At lesser depths and during other seasons, only a few individuals of this species were caught in the same area. This indicated a seasonal variability in local abundance that is most probably related to reproduction. Mora moro has an excellent flesh and may represent a valuable fisheries resource. Open problems requiring further research in the study area such as selectivity effects of the fishing gear and possible influences of ecological factors on geographical distribution of the deep-water ichthyofauna, as well as important aspects of the future demersal fisheries management in the Canarian archipelago are briefly outlined.