Phylogeography and genetic structure of the Canarian common chaffinch (Fringilla coelebs) inferred with mtDNA and microsatellite loci

The widespread common chaffinch (Fringilla coelebs) inhabits five of the seven Canary Islands. Sequences of the mitochondrial cytochrome b gene (1002 bp) revealed new insights into the systematics and phylogeography of this taxon. Additionally, a set of microsatellite loci were analyzed to examine the structure of these populations. Our results suggest that a new species of the genus Fringilla is present in the Canary Islands, which comprises at least three subspecies, but with a different distribution to that which has been morphologically accepted. The specimens from Gran Canaria are genetically distinct from those of La Gomera and Tenerife (F. c. canariensis), which suggests the existence of an undescribed taxon. Furthermore, nuclear microsatellite data suggest an ongoing incipient speciation process in this population. This study provides both important conservationist implications and a basis for re-evaluating the taxonomic status of the Canarian Fringilla coelebs populations.     

Changes in leaf nutrient traits in a wildfire chronosequence

The effect of wildfire on ecosystem function is gaining interest since climate change is expected to increase fire frequency and intensity in many forest systems. Fire alters the nutritional status of forest ecosystems, affecting ecosystem function and productivity, but further studies evaluating changes in leaf nutrient traits induced by forest wildfires are still needed. We used a 17-year-old Pinus canariensis wildfire chronosequence to elucidate the nature of nutrient limitations in natural and unmanaged pine forest in the Canary Islands. Pine needles were sampled in winter and spring and analysed for N and P concentrations. As expected, we found the lowest leaf N and leaf P in recently burned plots. However, the leaf N:P ratio was higher in burned versus unburned plots, suggesting that the decrease in P availability due to the fire is larger than that of N. For all leaf traits and sampling dates, leaf trait values in burned plots matched those observed in unburned plots 17 years after a fire. The N:P ratio found in P. canariensis needles was one of the lowest values reported in the literature for woody species, and suggests that all pine trees in the chronosequence are unambiguously limited by low N availability. Our results show that these N-limited pine forests retained N more efficiently than P 4 years after a wildfire; however, leaf N recovery is slower than P recovery, suggesting that the mechanisms responsible for pine N limitation operate continuously in these forests.     

Spatial pattern and variability in soil N and P availability under the influence of two dominant species in a pine forest

The presence of a legume in a nitrogen (N)-limited forest ecosystem may not only create ??islands of N fertility?? but also affect the phosphorus (P) availability. The main objective of this study was to compare the effect of a pine (Pinus canariensis) and a leguminous (Adenocarpus viscosus) species on the spatial pattern and variability of different labile organic-N (microbial biomass-N [MB-N] and dissolved organic-N [DON]), as well as inorganic-N (IN) and ?CP fractions (NH₄-N, NO₃-N, and PO₄-P), in a forest soil of the Canary Islands (Spain). Assuming some litter quantity and quality differences between these two species, we expected to find higher soil labile organic-N concentrations under isolated individuals of P. canariensis than under isolated individuals of A. viscosus. We also expected to find higher concentrations and spatial dependence (percentage of total variance explained by spatial autocorrelation) of NO₃-N beneath A. viscosus than beneath P. canariensis canopies, and higher spatial scaling of soil variables under the influence of P. canariensis canopies than under the presence of A. viscosus individuals. Moreover, we tested whether the soil variables measured under isolated individuals of both species showed a different spatial variability than the same soil variables measured under overlapping pine canopies inside a pine forest. To test these hypotheses, soil samples under isolated mature individuals of each species were collected in the winter and summer, whereas under a pine forest canopy, the sampling was performed only in the winter. The winter MB-N and DON concentrations were significantly higher beneath the pine individual, whereas the winter NO₃-N, NO₃-N-to-IN ratio, and PO₄-P were significantly higher under the leguminous individual; these differences were not observed in the summer samples. We found higher spatial ranges under the pine than under the legume canopy in the winter sampling, and the spatial dependence of NO₃-N was twice as high beneath the legumes as under the pines at both sampling dates. The soil spatial variability was higher (up to 17 times higher) under isolated individuals than inside the pine forest. The results of this study suggest that both the morphological and physiological characteristics of P. canariensis and A. viscosus, as well as the spatial pattern of P. canariensis, may influence the spatial pattern and variability of soil resources.     

Deep-water macroalgae from the Canary Islands: new records and biogeographical relationships

Due to the geographical location and paleobiogeography of the Canary Islands, the seaweed flora contains macroalgae with different distributional patterns. In this contribution, the biogeographical relations of several new records of deep-water macroalgae recently collected around the Canarian archipelago are discussed. These areBryopsidella neglecta (Berthold) Rietema,Discosporangium mesarthrocarpum (Meneghini) Hauck,Hincksia onslowensis (Amsler et Kapraun) P. C. Silva,Syringoderma floridana Henry,Peyssonnelia harveyana J. Agardh,Cryptonemia seminervis (C. Agardh) J. Agardh,Botryocladia wynnei Ballantine,Gloiocladia blomquistii (Searles) R. E. Norris,Halichrysis peltata (W. R. Taylor) P. Huvé et H. Huvé,Leptofauchea brasiliensis Joly, andSarcodiotheca divaricata W. R. Taylor. These new records, especially those in the Florideophyceae, support the strong affinity of the Canary Islands seaweed flora with the warm-temperate Mediterranean-Atlantic region. Some species are recorded for the first time from the east coast of the Atlantic Ocean, enhancing the biogeographic relations of the Canarian marine flora with that of the western Atlantic regions.     

Contrasting effects of wildfire and climate on radial growth of Pinus canariensis on windward and leeward slopes on Tenerife, Canary Islands

Little is known concerning the effects of wildfires on tree radial growth and their climatic response under contrasting regimes of fog water inputs on oceanic islands. On Tenerife, Canary Islands, windward slopes are humid with high-fog frequency due to influence of wet trade winds, while climate on leeward slopes is more arid. We used tree-ring records of Pinus canariensis Sweet ex Spreng. to quantify the effects of a fire of known date on radial growth and determine the main limiting climatic factors for growth. Radial growth patterns and their responsiveness to fire severity and climatic variation differed between windward and leeward slopes. Surface fire did not significantly impact growth, while crown fire caused short-term growth reduction, and even cessation, more pronounced on the windward slope. Growth rates, tree-ring common signal, and climate sensitivity were smaller on the windward slope, with cold winters, and summer water stress limiting growth. On the leeward slope, climate explained a greater amount of growth variation mainly due to negative effects of high October–December sea-level pressures causing dry winter conditions. Contrasting growth dynamics on both slopes may result from diverging physiological effects of water inputs and reduced radiation caused by fog drip. Our findings suggest that dating growth suppressions and absent rings are useful to date past high-severity crown fires in P. canariensis forests, in addition to ordinary fire scars dating indicative of low-severity surface fires.     

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     

Radiation of Atlantic goldcrests Regulus regulus spp.: evidence of a new taxon from the Canary Islands

Phylogenetic relationships between goldcrest populations from the Atlantic Islands (Azores and Canary Islands) were investigated by two molecular markers (mitochondrial control region and cytochrome b sequences), and partly by morphology and territorial song. The Azorean goldcrest populations are closely related to European nominate R. r. regulus. Most probably, the Azores were colonized by goldcrests in a single late‐pleistocene invasion, while colonization of the Canary Islands presumably occurred in two steps: An early invasion to Tenerife and La Gomera 1.9–2.3 million years (my) ago and a more recent one to El Hierro and La Palma 1.3–1.8 my ago. Distribution of haplotypes on the Azores suggests a division of R. r. azoricus on São Miguel into an eastern population with close affinities to R. r. sanctaemariae and a western population belonging to the lineage of R. r. inermis on the central and western island group. The Canarian populations are genetically substructured into a northeastern group embracing Tenerife and La Gomera and a second, southwestern group including El Hierro and La Palma. Genetic distances between members of the two Canarian clades range at 3.1–3.4% (TrN distance, control region and cytochrome b). Differentiation between the two groups is also supported by morphology and by territorial song. Substitution rate estimates for the both genes range at approximately the same values of 0.0031 and 0.0044 substitutions per site and lineage per my which roughly corresponds 0.61–0.83% divergence between Regulus lineages per my. Highest local rates occur in island clades of the Azorean and the Canarian population and in R. r. japonensis from the Russian Far East and Japan. However, a general acceleration of a molecular clock in island populations is not evident from the Regulus data set due to extremely low local rate estimates in the Canarian clade of Tenerife and La Gomera. As a taxonomic consequence of the marked differentiation of the two Canarian goldcrest clades the populations from El Hierro and La Palma are described as a taxon new to science and are named Regulus regulus ellenthalerae n. ssp.     

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.     

Factors controlling soil development in sand dunes: evidence from a coastal dune soil chronosequence

Aerial photographs, maps and optically stimulated luminescence dates were combined with existing soil data to construct high resolution chronosequences of soil development over 140 years at a temperate Atlantic UK dune system. Since soil formation had progressed for varying duration under different climate and nitrogen deposition regimes, it was possible to infer their relative influence on soil development compared with location-specific variables such as soil pH, slope and distance to the sea. Results suggest that soil development followed a sigmoid curve. Soil development was faster in wet than in dry dune habitats. In dry dunes, rates were greater than in the literature: they increased with increasing temperature and nitrogen deposition and decreased with increasing summer gales. The combination explained 62% of the variation. Co-correlation meant that effects of nitrogen deposition could not be differentiated from temperature. In wet dune habitats rates increased with temperature and decreased with gales. The combination explained only 23% of the variation; surprisingly, rainfall was not significant. Effects of location-specific variables were not significant in either habitat type. Nitrogen accumulation was faster in wet than dry dune habitats, averaging 43 kg N ha^?1 per year overall. Nitrogen accumulation greatly exceeded inputs from atmospheric deposition, suggesting rates of input for biological N fixation are 10–60 kg N ha^?1 per year. Recent climate and/or nitrogen deposition regimes may have accelerated soil development compared with past rates. These data suggest the importance of changing climate on soil development rates and highlight the contribution of biological N fixation in early successional systems.     

Clitopilus canariensis (Basidiomycota, Entolomataceae), a new species in the C. nitellinus-complex (Clitopilus subg. Rhodophana) from the Canary Islands (Spain)

Clitopilus canariensis is described as a new species belonging to Clitopilus subgenus Rhodophana on the basis of a collection from La Palma, Canary Islands. The new species grows on humus of Cistus monspeliensis and is known from a single site. A photograph of fresh basidiomes of the type-collection and line drawings are provided.     

Data on the family Pandalidae around the Canary Islands, with first record of Plesionika antigai (Caridea)

Of the 20 pandalid shrimps species and subspecies reported for the Eastern Central Atlantic (26–36° N), 16 were found in one or more Macaronesian archipelagos (Azores, Madeira, Canary Islands and Cape Verde Islands) (14–40° N), and 11 of them were recorded to date in the Canary Island waters (27° 30–29° 30 N): Bitias stocki Fransen, 1990; Heterocarpus ensifer ensifer A. Milne-Edwards, 1881; Heterocarpus grimaldii A. Milne-Edwards & Bouvier, 1900; Heterocarpus laevigatus Bate, 1888; Plesionika edwardsii (Brandt, 1851); Plesionika ensis (A. Milne-Edwards, 1881); Plesionika holthuisi Crosnier & Forest, 1968; Plesionika martia martia (A. Milne-Edwards, 1883); Plesionika narval (J.C. Fabricius, 1787); Plesionika williamsi Forest, 1964; and Stylopandalus richardi Coutière, 1905. In the present work, Plesionika antigai Zariquiey Álvarez, 1955 is recorded for the first time from the Canary Islands. As a result of many fishing surveys around the Canary Islands at 27–1550 m depth between 1985 and 1998, information on bathymetric distribution, habitat, size and biology of the 12 Canarian pandalid species is given. The geomorphologic, geographic and oceanographic characteristics of the Canary Islands marine ecosystems could explain the great diversity in the biogeographic patterns of the pandalid species inhabiting this area. The distribution patterns found were: Macaronesian (1 spec.), Atlanto-Mediterranean (1 spec.), Eastern Atlantic warm-temperate (1 spec.), amphi-Atlantic warm (2 spec.), amphi-Atlantic warm-temperate (1 spec.), pantropical (5 spec.), and cosmopolitan (1 spec.).     

Microsatellite loci isolation in the endangered Gran Canarian blue chaffinch (Fringilla teydea polatzeki) and their utility in closely related taxa

The blue chaffinch, Fringilla teydea, is an endemic species of the Canary Islands. This species is formed by two subspecies: The Teneriffean blue chaffinch (F. t. teydea), and the endangered Gran Canarian blue chaffinch, (F. t. polatzeki). Here we report the isolation and characterization of nine tetranucleotide microsatellites (AAAG and AAAT) from the Gran Canarian subspecies, using an enrichment protocol. An average of 7.8 alleles per locus and an average observed heterozygosity of 0.773 were found (n = 28). The loci were tested for their ability to cross amplify in the Teneriffean subspecies and in the common chaffinch (Fringilla coelebs). These microsatellites will be used to manage a captive breeding programme for the endangered Gran Canarian subspecies.     

Gas exchange characteristics of<Emphasis Type="Italic"> Pinus canariensis</Emphasis> needles in a forest stand on Tenerife,Canary Islands

Gas exchange characteristics and chlorophyll fluorescence of the Canarian endemic pine ( Pinus canariensis) were measured during the day for a year in a field stand on Tenerife, Canary Islands, Spain. Diurnal tendencies of gas exchange were variable depending on ambient conditions. In general they paralleled photosynthetic photon flux density with only one peak at midday, except on summer days with high air vapour pressure deficit (VPD), when needles exhibited a severe midday depression of CO ₂ assimilation rate ( A), in parallel with a reduction of stomatal conductance ( g _s). The internal CO ₂ concentration tendencies during the day suggest that stomatal closure was the main cause of the midday depression of photosynthesis. Chlorophyll fluorescence data corroborate this assertion, with the parameter F _v/ F _m reaching high values throughout day and year. P. canariensis living in the sub-tropic exhibited high values of A (maximal A value of 17 µmol m ^-2 s ^-1) and high optimal needle temperature for photosynthesis (25°C) which were at the upper limit of the values given for conifers and similar only to data obtained for some pine species adapted to habitats at similar latitudes. g _s was reduced to half when VPD attained 40 mbar, allowing this pine to have high A/ g _s values during high evaporative demand conditions.     

DNA-based confirmation that the parasitic wasp <Emphasis Type="Italic">Cotesia glomerata</Emphasis> (Braconidae,Hymenoptera) is a new threat to endemic butterflies of the Canary Islands

Island-endemic species can be particularly vulnerable to alien invasion. There are many examples of introduced insect parasitoids having a serious impact on endemic butterflies and moths. In 2006, a population of parasitic wasps was reared from larvae of the Canary Island Large White butterfly (Pieris cheiranthi), an endemic inhabitant of laurel forests unique to the Canary Islands of Macaronesia. Parasitoids were tentatively identified as Cotesia glomerata (Braconidae, Hymenoptera), a widely introduced agricultural bioagent. To corroborate this finding we sequenced 632 bp of mitochondrial cox1 from parasitoids and hosts from La Palma and from the native range of C. glomerata in continental Europe. These were combined with GenBank sequences and a character-based, phylogenetic approach was used to assess the species status of parasites and hosts. The La Palma parasitoid could unambiguously be assigned to C. glomerata under the criterion of diagnosibility with corroboration from multiple lines of evidence (DNA, morphology). We suggest that this opportunistic, non-native parasitoid wasp will be a threat to P. cheiranthi and other endemic Canarian butterflies. Parasitoid populations were recorded from P. cheiranthi in marginal forest habitats but not in central forest areas, suggesting that comprehensive habitat conservation of the Canarian laurel forests could prevent penetration of the alien parasitoid wasps and subsequent mortality of endemic butterfly populations.     

Water availability drives stem growth and stem water deficit of <Emphasis Type="Italic">Pinus canariensis</Emphasis> in a drought-induced treeline in Tenerife

Tree growth of Pinus canariensis at treeline in Tenerife, Canary Islands, is thought to be primarily controlled by wet season precipitation (P) prior to the current year´s growth. Therefore, we investigated the inter-annual variations in stem water deficit (ΔW) and radial growth (RG) during two consecutive years differing in wet season P. ΔW was extracted from stem circumference variations, and the influence of environmental variables was evaluated by Pearson correlation statistics. Wet season P was considerably lower in 2008 than in 2009; despite this difference in P between both years, shallow soil water availability was almost exhausted during both summers. However, the effect of shallow soil drought showing a clear seasonality of ΔW and RG was only detected in 2008. In summer 2009, RG rates were highest during the summer indicating that P. canariensis was able to tap water from deep soil layers originating from P prior to the current year´s growth. The ability to use deep soil water during extended periods of shallow soil water deficit was also reflected in a close positive correlation between RG and whole-tree water use. In our study, the effect of only one hydrological dry year resulted in a severe reduction in annual RG. Thus, when wet season P is low for a number of years, chronic drought may have negative implications for tree growth at treeline in Tenerife.     

Linking chronosequences with the rest of the world: predicting soil phosphorus content in denuding landscapes

Results from chronosequences from the arctic to the tropics show that phosphorus (P) availability, total P, and the fraction of bedrock-derived P remaining in soil diminishes as soils age. Thus we predict that ecosystems mantling old substrates are likely to have low available P. Yet there are myriad examples in the biogeochemical literature where the results from chronosequences are used to argue the reverse, and ecosystems observed to be P poor are assumed to mantle an old substrate. This premise is difficult to test, for while the concept of substrate age is useful on uneroded surfaces that formed at a particular time, it becomes obscured in denuding landscapes, where substrate ages instead reflect the rates of rock weathering, denudation and mixing of dust into soil. Here we explore this premise as it relates to one of the most ubiquitous assumptions in the biogeochemical literature: that the differences in nitrogen (N) and P cycling between temperate and tropical regions are driven by gradients in substrate age. We build a conceptual framework for quantifying the fraction of parent material P remaining in soil ([SoilP]/[RockP]), by estimating P inputs (rock weathering and dust deposition) and outputs (P leaching). We parameterize our model with spatially explicit (0.5°) estimates of global denudation, weathering zone thickness, and P deposition. To test the assumption that latitudinal gradients in P status are the result of soil age, we apply a single P loss rate, derived from a humid tropical system in the Hawaiian Islands, to our spatially explicit map of soil residence times. Surprisingly, in this formulation, we find only a modest latitudinal gradient in soil P depletion, with mean depletion values in the humid tropics <2× greater than in the previously unglaciated humid temperate zone. This small latitudinal gradient in P depletion is unlikely to be sufficient to drive the observed differences in tropical vs. temperate ecosystem stoichiometry (e.g. trends in foliar and litter N:P). Thus our results suggest that, to the extent P depletion is greater in the tropics, the appropriate conceptual model for attributing causation may not be one of a chronosequence where time is the primary driver of P loss. We hypothesize that the covariation of inferred P availability with latitude may be strongly controlled by latitudinal changes in rates of P leaching and occlusion, rather than gradients in substrate age.     

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.     

Exploring the effect of host tree identity on epiphyte bryophyte communities in different Canarian subtropical cloud forests

This study explores the host tree preferences of epiphyte bryophyte communities in two key types of evergreen cloud forests on La Gomera (Canary Islands, Spain): ericaceous versus broadleaved laurel forest. By comparing two pairs of tree species (Erica arborea and Myrica faya vs. Ilex canariensis and Laurus novocanariensis) that occur in both forest types, we quantitatively examined whether epiphyte-host relationships change with the type of forest. In 51 ancient forest stands, the low-trunk bryophyte composition on the two host tree groups was analysed with both non-parametric procedures and ordination analyses. Our results highlight that the influence of local environmental conditions on the structuring of epiphyte communities may be different depending on the host tree identity. Whilst the epiphyte composition of the host tree group formed by I. canariensis and L. novocanariensis differed significantly between forest types depending on solar exposure, we failed to find variations for the other host tree group, excepting in those localities wherein mist precipitation was extremely high. Our findings highlight the importance of the natural diversity of tree species along environmental gradients, as well as of water availability at different spatial scales for epiphytic bryophytes in montane cloud forests.     

Horizontal gene transfer from a flowering plant to the insular pine Pinus canariensis (Chr. Sm. Ex DC in Buch)

Horizontal gene transfer (HGT) is viewed as very common in the plant mitochondrial (mt) genome, but, to date, only one case of HGT has been found in gymnosperms. Here we report a new case of HGT, in which a mt nad5-1 fragment was transferred from an angiosperm to Pinus canariensis. Quantitative assay and sequence analyses showed that the foreign nad5-1 is located in the mt genome of P. canariensis and is nonfunctional. An extensive survey in the genus Pinus revealed that the angiosperm-derived nad5-1 is restricted to P. canariensis and present across the species'' range. Molecular dating based on chloroplast DNA suggested that the HGT event occurred in the late Miocene after P. canariensis split from its closest relatives, and that the foreign copy became fixed in P. canariensis owing to drift during its colonization of the Canary Islands. The mechanism of this HGT is unclear but it was probably achieved through either direct cell–cell contact or external vectors. Our discovery provides evidence for an important role of HGT in plant mt genome evolution.