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.     

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.     

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.     

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.     

Phylogeography of skinks (Chalcides) in the Canary Islands inferred from mitochondrial DNA sequences

Mitochondrial DNA (mtDNA) evolution was investigated in skinks of the genus Chalcides found in the Canary Islands ( Ch. sexlineatus, Ch. viridanus and Ch. simonyi ), together with some North African congenerics ( Ch. polylepis and Ch. mionecton ). Several sites were included within islands to cover areas of known within-island geographical variation in morphology. Skinks from the islands of El Hierro and La Gomera appear to be sister taxa. The relationships between this clade and the Tenerife and Gran Canarian skinks were not fully resolved, although the best working hypothesis indicated monophyly with the former, with the latter forming a closely related outgroup. Ch. simonyi from Fuerteventura was more distantly related to the Western Canary Island skinks and did not show close relationships with the North African species Ch. mionecton and Ch . polylepis . Possible colonization sequences for the four most Western Canary Islands were considered. El Hierro appears to have been colonized relatively recently from La Gomera, commensurate with the recent origin of this island, while dispersal between La Gomera and Tenerife and between Gran Canaria and Tenerife or La Gomera appears to have taken place considerably earlier. Substantial within-island haplotype divergence was found in Gran Canaria and Tenerife. This may be a result of recent periods of intense volcanic activity found within these two islands. Lower levels of within-island differentiation are found in La Gomera and El Hierro and may be explained by lower levels of volcanic activity during recent geological history and a more recent colonization, respectively.     

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.     

Raptors as legitimate secondary dispersers of weed seeds

Although raptors are not usually seen as seed dispersers, we demonstrate their important ecological role as legitimate seed dispersers of weeds. On the oceanic island of Fuerteventura in the Canary Islands, up to 11 000 weed seeds from a total of 39 and 62 different plant species, respectively, were found in 300 and 319 pellets, respectively, of two native predatory birds, the Common Buzzard Buteo buteo and the Eurasian Kestrel Falco tinnunculus, which consumed European Rabbits Oryctolagus cuniculus and Barbary Ground Squirrels Atlantoxerus getulus, both invasive mammals that carried these seeds in their guts. Four and seven weed species appeared with a frequency of occurrence greater than 10% in the pellets of Kestrels and Buzzards, respectively. Despite the time interval between pellet collection and germination experiments, almost 10% of planted seeds germinated, even after passing through two digestive processes, one inside the mammal and another inside the raptor. Our study encourages a change of perspective in this regard, as raptors may be involved in complex interactive dispersal processes on a wider geographical scale.     

Microhabitat selection of the European rabbit on La Palma, Canary Islands, Spain

Little information exists regarding habitat selection on oceanic islands for the introduced European rabbitOryctolagus cuniculus Linnaeus, 1758. I employed the sharp altitude gradient of one of the Canary Islands, La Palma, to analyze factors relating to rabbit abundance. I used pellet counts to assess rabbit abundance in relation to microhabitat variables during June and July of 1999 and 2001. Number of rabbits varied with grass cover in the different biotopes for both years. They were particularly abundant in biotopes at low elevation and with rich grass cover. Rabbits were more numerous in 2001 than in 1999, and selected shrub protection in places with high levels of grass cover. Areas with vegetation transformed by humans were particularly suitable for wild rabbits, in contrast to less disturbed island biotopes. Rabbit abundance was usually associated with rich grass cover, which might be a limiting factor on La Palma.     

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.     

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.     

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.     

Diversification of the forest beetle genus Tarphius on the Canary Islands, and the evolutionary origins of island endemics

The flightless beetle genus Tarphius Erichson (Coleoptera: Colydiidae) is a distinctive element of the beetle fauna of the Canary Islands with 29 species distributed across the five western islands. The majority of Tarphius species are rare and intimately associated with the monteverde forest and only two species occur on more than one island. In this study we investigate the phylogeography of the Canary Island Tarphius, and their relationship to Tarphius from the more northerly archipelagos of Madeira and the Azores using maximum parsimony and Bayesian inference analysis of mitochondrial cytochrome oxidase I and II sequence data. We use geological datings for the Canary Islands, Azores, and Madeira to calibrate specific nodes of the tree for the estimation of divergence times using a penalized likelihood method. Data suggest that the Canary Island species assemblage is of some antiquity, however, much of this species diversity is relatively recent in origin. The phylogenetic relationships of species inhabiting the younger islands of El Hierro and La Palma indicate that colonization events between islands have probably been a significant factor in the evolutionary history of the Canary Island species assemblage. A comparison of molecular phylogenetic studies of arthropods on the Canary Islands suggests that, in the evolution of the arthropod species community of an island, the origin of endemic species is initially the result of colonizing lineages differentiating from their source populations. However, as an island matures a greater proportion of endemic species originate from intra-island speciation.     

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.     

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.     

Reproduction in an endemic bird of a semiarid island: a food-mediated process

We analyse the effects of rainfall, temperature, food availability and nest predation on the between-year variation in reproduction of the Canary Islands stonechat Saxicola dacotiae , a bird species endemic of the semiarid island of Fuerteventura (Canary Islands, Spain). We monitored the reproductive performance and output of the species across its whole distribution area during three consecutive breeding periods, also measuring rainfall, temperature and food (arthropod) availability on the ground monthly. Rainfall varied from 27.3 mm in 2000–2001 (dry year) to 124.5 mm and 125.1 mm (average years), respectively, in the 2001–2002 and 2002–2003 breeding seasons. The onset of breeding closely matched variation in the onset of the autumn-winter rains among years. Arthropod availability was strongly and positively related to rainfall one month before. Reproductive investment (number of clutches and clutch size) was correlated to arthropod availability both among years and among sites within years. Stonechat pairs bred once or did not breed at all in the dry year whereas they bred twice in the other two years. Clutch size was smaller in the dry year and larger in the second as compared with the first in the other two years. Reproductive investment largely determined reproductive output (number of fledglings) as there were no significant spatial or temporal variation in hatching success (90% on average) or nest predation (29% on average, mostly due to feral cats Felis catus ). Within- and among years variability in temperature or predation did not match variability in the onset, length or reproductive investment and output recorded throughout the study years. These results suggest that variability in annual fecundity in the Canary Islands Stonechat was mainly driven by rainfall through a food-mediated process.     

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.     

How important are pelagic preys for the kelp gull during chick-rearing at the South Shetland Islands?

An analysis of pellets regurgitated indicated adult kelp gulls (Larus dominicanus) on the South Shetland Islands consumed predominantly intertidal prey, whereas previous studies at Antarctic Peninsula sites have reported kelp gulls consuming predominantly pelagic species. The pellets collected at Nelson Island during the chick-rearing period indicated that the limpet Nacella concinna was their most frequent prey, followed by carrion, gammariids, snails and krill. Fish were scarcely represented. Also, regurgitated stomach contents of chicks showed that limpets and carrion were the most frequent food items, accounting for 70% of the mass. However, gammariids were particularly important by number. Significant differences were observed in the overall comparison of the diet as reflected by the two sampling methods. In general, the importance of pelagic prey was negligible when compared to intertidal or scavenged prey. Our results differ greatly from those reported for the Antarctic Peninsula, where chicks were almost exclusively fed with the pelagic fish Pleuragramma antarcticum. These differences could be related to the abundance of pelagic resources in southernmost latitudes, and/or to the presence of more extensive intertidal foraging areas at the South Shetland Islands. Received: 7 October 1996 / Accepted: 14 July 1997     

Evidence of connectivity between continental and differentiated insular populations in a highly mobile species

Aim Genetically differentiated insular populations are candidates for independent units for conservation. However, occasional immigration to reduced island populations may occur and potentially have important consequences in their future viability and evolutionary potential. In this study, we investigate the conservation implications of population structure and connectivity of insular and continental populations of a migratory raptor as determined using genetic tools and satellite tracking. Location Western European populations in the Iberian Peninsula and two insular populations in the Mediterranean Sea (Balearic Islands) and Atlantic Ocean (Canary Islands). Methods We genotyped 22 microsatellite loci in 96 Egyptian vultures (Neophron percnopterus) from the Iberian Peninsula, 36 from Menorca (Balearic archipelago) and 242 (85% of the current population) from Fuerteventura (Canary Islands). We analysed genetic variation to estimate structure, gene flow, genetic diversity, effective size and recent demographic history of the populations. Additionally, 19 vultures were marked with satellite transmitters to track their migration routes. Results Insular populations were genetically differentiated from those of the mainland. We detected immigration in the insular populations and within the continental counterpart. We found similar levels of genetic variability between the continent and the islands, and a bottleneck analysis indicated recent sharp population declines in both archipelagos but not on the continent. Main conclusions Our study provides evidence that, in spite of significant differentiation, insular populations of highly mobile species may remain connected with the mainland. Conservation programmes should take into account population connectivity and integrate differentiated units of management within complex units of conservation that can best maintain processes and potential for evolutionary change.