Out of Africa: north‐westwards Pleistocene expansions of the heather Erica arborea

Aim The heather Erica arborea L. is a dominant element of the circum‐Mediterranean region. Its broad, disjunct distribution, ranging from Macaronesia to eastern Africa, is consistent with the fragmentation of the evergreen tropical and subtropical forests that dominated Europe and North Africa in the Tertiary. This study aims to investigate phylogeographical patterns in E. arborea and to determine whether the current disjunct distribution of the species is a relict of a once wider distribution, or a recent range expansion in response to the establishment of suitable conditions. Location Mediterranean, Macaronesia, North and eastern Africa. Methods A total of 105 samples were collected across the species’ distribution range and sequenced at four cpDNA loci (atpB–rbcL, matK, trnH–psbA and rpl16). Phylogenetic reconstructions, molecular dating techniques and Bayesian ancestral area reconstructions were used in combination with population genetic statistics (haplotype diversity, N_ST, F_ST, Fu’s F_S) to describe the pattern of present genetic diversity in E. arborea and infer its biogeographical history. Results Haplotype diversity in Macaronesia and the east and central Mediterranean is much lower than that observed in eastern Africa/Arabia and the western Mediterranean. Bayesian ancestral area reconstructions and molecular dating suggest that E. arborea colonized the Mediterranean westwards from eastern Africa/Arabia at least twice during a time period ranging between the upper Miocene and the upper Pleistocene. Main conclusions The phylogeography of E. arborea involves a complex history of range expansions and contractions, which has resulted in a pattern of distribution that mimics that expected for a Tertiary vicariance event. Despite the presence of a late Tertiary refugium in the Iberian Peninsula, the current distribution of the species throughout the Mediterranean is explained by a Pleistocene expansion originating from eastern Africa. One explanation for the isolation of the Iberian refugium is the rapidity of the most recently identified colonization wave, as inferred by the absence of global phylogeographical signal in the data and significantly negative values of Fu’s F_S statistic for European populations. Macaronesia was colonized during each of these two expansion waves, confirming that the laurisilva (laurel forest flora) is a complex entity including both ancient relicts and recent colonizers.     

Molecular,phytochemical and morphological characterization of the liverwort genus Radula in Portugal (mainland,Madeira, Azores)

Relationships of the eight species of the liverwort genus Radula occurring in Portugal (mainland, the Madeira and Azores archipelagos), including the Macaronesian endemics R. jonesii and R. wichurae, were evaluated based on molecular, phytochemical and morphological–anatomical data. Maximum parsimony and Bayesian analyses were performed with sequences from three plastid DNA markers (trnS _GGA-rps4 spacer, rps4 gene, trnL _UAA intron), volatile oil compounds, as well as qualitative morphological–anatomical characters. In addition, the molecular data were subjected to maximum likelihood analysis. The eight taxa, R. aquilegia, R. carringtonii, R. complanata, R. holtii, R. jonesii, R. lindenbergiana, R. nudicaulis and R. wichurae, can be clearly distinguished from each other, either by molecular data alone or by combination of characters from all three data sets. Radula aquilegia is monophyletic according to the molecular data, but shows considerable, yet undescribed intraspecific morphological and phytochemical variability. Recognition of R. complanata and R. lindenbergiana as separate species, previously based solely on the paroecious vs. dioecious sexual condition, is moderately supported by the molecular phylogenetic analyses and strongly supported by the phytochemical data. The Radula species, narrowly distributed in Macaronesia and Atlantic Europe, probably have two different origins. For Radula holtii and R. nudicaulis, connections with Radula species from the Neotropics are indicated. The other species, among them the two Macaronesian endemics, are closely related with the R. complanata/R. lindenbergiana complex, which is widely distributed in the northern hemisphere.     

Two genetic divergence centers revealed by chloroplastic DNA variation in populations of <Emphasis Type="Italic">Cinnamomum kanehirae</Emphasis> Hay

Cinnamomum kanehirae Hayata (Lauraceae), the most valuable subtropical and temperate broadleaf timber tree in Taiwan, is rapidly disappearing from the wild. Taking advantage of a scion garden established by the Taiwan Forestry Research Institute, we examined patterns of chloroplast DNA (cpDNA) variations in 19 populations including 94 individuals. By sequencing two cpDNA fragments using universal primers (the trnL-trnF and petG-trnP intergenic spacers), we found eight polymorphic sites, six haplotypes, and extremely low nucleotide diversity (π = 0.00016) from 792 bp aligned sequences. The ancestral haplotype is widely distributed. Among the populations studied, three separated populations, at Yungfeng, Fuli, and Tahu have high nucleotide diversity. No phylogeographical structures of haplotypes were revealed because the tests of N _ST−G _ST for populations did not differ from zero in any situations; a ‘star-like’ genealogy is characteristic when all haplotypes rapidly coalesce and is a general outcome of population expansion. The neutrality test also suggested demographic expansion. The genetic divergence and diversity analyses suggested that two potential refugia existed during the last glaciation with a major one located in southeastern Taiwan and a minor one located in Tahu in north-central Taiwan in the Hsuehshan Range, west of the Central Mountain Range.     

Phylogeographic analysis of the endemic species Sibiraea angustata reveals a marginal refugium in the Qinghai–Tibet Plateau

The phylogeography of Sibiraea angustata, an endemic shrub species, was studied in the Qinghai–Tibet plateau (QTP). We investigated 466 individuals of S. angustata from 39 populations basically covering its total distribution area. Eight haplotypes (A–H) were detected by sequencing the intergenic chloroplast spacer trnS–trnG (600 bp), and one ancestral haplotype (A) was found to be widely distributed. The level of differentiation among populations was very high (G_ST=0.768; N_ST=0.850) and a significant phylogeographical structure was revealed (N_ST>G_ST). Analysis of molecular variance (AMOVA) similarily revealed a high level of differentiation among populations (84%, F_ST=0.842), indicating that little gene flow has occurred among populations mutually isolated by high mountains and rivers in the QTP. On the QTP platform there was only one widespread haplotype (A) in most populations, while populations along the eastern and southeastern edges had high diversity and unique haplotypes. Our results suggest that a glacial refugium may have been located on the eastern or southeastern edges of QTP during the last glaciation, and that interglacial and postglacial range expansion occurred from that refugium. Nested clade analysis (NCA) also suggests this scenario, which indicates that the current spatial distribution of cpDNA haplotypes and populations mainly resulted from long distance colonization, possibly coupled with subsequent or past fragmentation followed by range expansion and allopatric fragmentation.     

Recent origin,active speciation and dispersal for the lichen genus Nephroma (Peltigerales) in Macaronesia

Aim We reconstructed the phylogeny of the lichen genus Nephroma (Peltigerales) to assess the relationships of species endemic to Macaronesia. We estimated dates of divergences to test the hypothesis that the species arose in Macaronesia (neo‐endemism) versus the oceanic archipelagos serving as refugia for formerly widespread taxa (palaeo‐endemism). Location Cosmopolitan with a special focus on the archipelagos of the Azores, Madeira and the Canary Islands. Methods DNA sequences were obtained from 18 species for three loci and analysed using maximum parsimony, maximum likelihood and Bayesian inferences. Divergence dates were estimated for the internal transcribed spacer (ITS)‐based phylogeny using a relaxed molecular clock. Reconstruction of the ancestral geographical range was conducted using the Bayesian 50% majority rule consensus tree under a parsimony method. Results The backbone phylogenetic tree was fully supported, with Nephroma plumbeum as sister to all other species. Four strongly supported clades were detected: the Nephroma helveticum, the N. bellum, the N. laevigatum and the N. parile clades. The latter two share a common ancestor and each includes a widespread Holarctic species (N. laevigatum and N. parile, respectively) and all species endemic to Macaronesia. The data suggest a neo‐endemic origin of Macaronesian taxa, a recent range expansion from Macaronesia of both widespread species, a range expansion limited to the Mediteranean Basin and south‐western Europe for another taxon, and a long dispersal event that resulted in a speciation event in the western parts of North America. Main conclusions The Macaronesian endemic species belong to two sister clades and originated from a most recent common ancestor (MRCA) shared with one widely distributed taxon, either N. parile or N. laevigatum. Estimates of the mean divergence dates suggest that the endemics originated in the archipelagos after the rise of the volcanic islands, along with the ancestor to the now widespread species, which probably expanded their range beyond Macaronesia via long‐distance dispersal. This study provides the first phylogenetic evidence of Macaronesian neo‐endemism in lichenized fungi and provides support for the hypothesis that oceanic islands may serve as a source for the colonization of continents. However, further data are needed to properly assess the alternative hypothesis, namely colonization from western North America.     

Disentangling the genetic and morphological structure of Patella candei complex in Macaronesia (NE Atlantic)

The uptake of natural living resources for human consumption has triggered serious changes in the balance of ecosystems. In the archipelagos of Macaronesia (NE Atlantic), limpets have been extensively exploited probably since islands were first colonized. This has led to profound consequences in the dynamics of rocky shore communities. The Patella candei complex includes various subspecies of limpets that are ascribed to a particular archipelago and has been the focus of several taxonomic surveys without much agreement. Under a conservational perspective, we apply morphometric and genetic analyses to test subspecies boundaries in P. candei and to evaluate its current population connectivity throughout Macaronesia (Azores, Madeira, and Canaries). A highly significant genetic break between archipelagos following isolation by distance was detected (F_ST = 0.369, p < .001). Contrastingly, significant genetic differentiation among islands (i.e., Azores) was absent possibly indicating ongoing gene flow via larval exchange between populations. Significant shell‐shape differences among archipelagos were also detected using both distance‐based and geometric morphometric analyses. Adaptive processes associated with niche differentiation and strong barriers to gene flow among archipelagos may be the mechanisms underlying P. candei diversification in Macaronesia. Under the very probable assumption that populations of P. candei from each archipelago are geographically and/or ecologically isolated populations, the various subspecies within the P. candei complex may be best thought of as true species using the denomination: P. candei in Selvagens, Patella gomesii in Azores, Patella ordinaria in Madeira, and Patella crenata for Canaries. This would be in agreement with stock delimitation and units of conservation of P. candei sensu latu along Macaronesia.     

The barriers to oceanic island radiation in bryophytes: insights from the phylogeography of the moss Grimmia montana

Aim In contrast to angiosperms, bryophytes do not appear to have radiated in Macaronesia and the western Mediterranean. We evaluate if: (1) the apparent lack of radiation in bryophytes reflects our failure to recognize cryptic endemic species; (2) bryophytes are characterized by extremely low evolutionary rates; or (3) bryophytes have a high dispersal ability, which prevents genetic isolation. Location Worldwide, with a special emphasis on Macaronesia and the western Mediterranean. Methods Three chloroplast regions were sequenced from samples of the moss Grimmia montana from its entire distribution range. Network analyses, Fst and Nst statistics were used to describe and interpret the phylogeographical signal in the data. Results Despite significant phylogeographical signal in the chloroplast genome, which demonstrates limits to gene flow at the continental scale, repeated sister group relationships observed among accessions from different geographical areas suggest recurrent colonization patterns. These observations are consistent with mounting evidence that intercontinental distributions exhibited by many bryophyte species result from long‐distance dispersal rather than continental drift. Madeiran and western Mediterranean island haplotypes are either shared by, or closely related to, European and North American ones. Fst values between Madeira, western Mediterranean islands, North America and Europe are not significantly different from zero, and suggest that Madeira and the south‐western Mediterranean are subject to strong transatlantic gene flow. By contrast, haplotypes found in the Canary Islands are shared or closely related to those of populations from south‐western Europe or southern Africa. Main conclusions Multiple origins and colonization events are not consistent with the hypothesis of a relictual origin of the Macaronesian moss flora. One possible reason for the failure of taxa that experienced multiple colonization events to radiate is niche pre‐emption. We suggest that strong gene flow, coupled with the occupancy of all suitable niches, either by earlier conspecific colonizers or by other species, could be the mechanism preventing island radiation in G. montana and other cryptogams with high long‐distance dispersal abilities.     

Mitochondrial DNA patterns in the Macaronesia islands: Variation within and among archipelagos

Macaronesia covers four Atlantic archipelagos: the Azores, Madeira, the Canary Islands, and the Cape Verde islands. When discovered by Europeans in the 15th century, only the Canaries were inhabited. Historical reports highlight the impact of Iberians on settlement in Macaronesia. Although important differences in their settlement are documented, its influence on their genetic structures and relationships has yet to be ascertained. In this study, the hypervariable region I (HVRI) sequence and coding region polymorphisms of mitochondrial DNA (mtDNA) in 623 individuals from the Azores (120) and Canary Islands (503) were analyzed. Combined with published data, these give a total of 1,542 haplotypes from Macaronesia and 1,067 from the Iberian Peninsula. The results obtained indicate that Cape Verde is the most distinctive archipelago, with an mtDNA pool composed almost exclusively of African lineages. However, the other archipelagos present an mtDNA profile dominated by the presence of West‐Eurasian mtDNA haplogroups with African lineages present in varying proportions. Moreover, no signs of integration of typical Canarian U6 lineages in the other archipelagos were detected. The four Macaronesia archipelagos currently have differentiated genetic profiles, and the Azores present the highest intra‐archipelago differentiation and the lowest values of diversity. The analyses performed show that the present‐day genetic profile of the Macaronesian archipelagos was mainly determined by the initial process of settlement and further microdifferentiation probably as a consequence of the small population size of some islands. Moreover, contacts between archipelagos seem to have had a low impact on the mtDNA genetic pool of each archipelago. Am J Phys Anthropol, 2010. © 2009 Wiley‐Liss, Inc.     

Phylogeographic analyses suggest that a deciduous species (Ostryopsis davidiana Decne., Betulaceae) survived in northern China during the Last Glacial Maximum

Aim Palaeontologial data suggest that all temperate forest species in northern China migrated southwards during the Last Glacial Maximum (LGM) and recolonized post‐glacially within the Holocene. We tested this assumption using phylogeographical studies of a temperate deciduous shrub species (Ostryopsis davidiana Decne., Betulaceae), which has a wide distribution in northern China. Location Northern China. Methods We sequenced two chloroplast DNA (cpDNA) fragments (trnL–trnF and psbA–trnH, together about 1300 bp in length) of 294 plants from 21 populations across the total distribution range of this species. We used maximum parsimony and haplotype network methods to construct phylogenetic relationships among haplotypes. Results The analysis of cpDNA variation identified eight haplotypes. A single haplotype was fixed in all populations except for one population that was polymorphic, having two haplotypes. The population subdivisions were extremely high (G_ST = 0.972 and N_ST = 0.974), suggesting very low gene flow between populations. Haplotypes clustered into two tentative clades, both of which occur in the southern region of the species’ range but only one of which occurs in the northern region. Across the sampled populations, the haplotype distributions were differentiated geographically. Main conclusions Our analyses suggest that multiple refugia were maintained across the range of O. davidiana in both northern (north of the Qing Mountains) and southern (south of the Qing Mountains) regions during the LGM rather than that the species survived only in the south and subsequently colonized northwards. The extremely low within‐population diversity of this species suggests strong bottleneck or founder effects within each fragmented region during the Quaternary climatic oscillations. These findings provide important clues for understanding range shifts and changes in within‐ and/or between‐population genetic diversity of temperate forests in response to past climatic oscillations in northern China.     

青藏高原濒危植物唐古特大黄的遗传多样性

唐古特大黄(Rheum tanguticum)是中国传统的中藏药材,近几年由于生境的严重破坏,已濒临灭绝,并被列入濒危植物名单。为了探索唐古特大黄物种濒危的原因并保护其野生资源,本研究采集了9个居群87个个体的唐古特大黄样本,基于该物种的叶绿体基因trn S-G序列对其进行了遗传多样性研究。结果表明,唐古特大黄物种具有较高的遗传多样性水平(Ht=0.694),其中95.97%的遗传分化来自于居群间(G_(ST)=0.960),4.03%的遗传分化来自于居群内(Hs=0.028)。AMOVA分析也显示唐古特大黄居群间基因流较小(N_m=0.01),存在较高的遗传分化(F_(ST)=0.9631)。唐古特大黄较高的遗传多样性水平可能与该物种较长的进化史和生活史有关,居群间较高的遗传分化可能与高山地区特殊的地理环境和人类活动有关。根据研究结果,建议对唐古特大黄所有野生居群进行就地保护,同时收集种质资源开展异地繁殖工作,以保护物种的遗传多样性,维持其进化潜力。     

Biogeographic patterns in Mediterranean and Macaronesian species of Saxifraga (Saxifragaceae) inferred from phylogenetic analyses of ITS sequences

A biogeographic study of Saxifraga section Saxifraga was performed based on phylogenetic analyses of ITS (internal transcribed spacer) sequences of nuclear ribosomal DNA. ITS sequences from 21 species and 31 populations were examined to identify colonization patterns for the two species of Saxifraga occurring in Macaronesia and for S. globulifera in the west Mediterranean basin. Phylogenetic analysis of the sequence data yield a single most parsimonious tree with many of the major clades well supported by bootstrap and decay values. The ITS tree provided resolution at specific and populational levels that points to two biogeographic patterns within the genus. In contrast to the molecular evidence provided by other authors for a Mediterranean origin of several Macaronesian genera of angiosperms, our results indicate that the Madeiran archipelago was colonized a single time by a species of Saxifraga originating from the Eurosiberian region. On the other hand, the molecular evidence also suggests that populations of S. globulifera from North Africa have been isolated for a long time from populations occurring in the Iberian Peninsula, and that the endemic S. reuteriana has evolved from the Iberian populations of S. globulifera. The Mediterranean Sea has probably been an effective isolating barrier for some plant groups that occur in Europe and North Africa.     

Clonal diversity and geographic structure in <Emphasis Type="Italic">Pleurochaete squarrosa</Emphasis> (Pottiaceae): different sampling scale approach

This paper describes our investigation of genetic variation and clonal structure of the Mediterranean moss Pleurochaete squarrosa (Brid.) Lindb. (Pottiaceae), using inter-simple sequence repeat (ISSR) molecular markers and trnL_UAA (intron of plastid gene for Leu tRNA) sequence, choosing different sampling strategies and scales on 16 European populations. Moreover, the intercontinental distribution of two trnL haplotypes, previously found over a large area and including 24 populations in three continents, was also investigated. Despite the prevalent asexual reproduction, P. squarrosa shows a high level of genetic diversity. Some site features seem to affect the clonal structure at the local scale, influencing the relocation of detached fragments and the level of intermingling, but they do not substantially affect genetic diversity. The peculiar vegetative reproduction coupled with somatic mutation could partly account for the genetic variation detected. Genetic distances highlight geographic isolation and limited gene flow among populations. We found only two trnL haplotypes in Europe due to length polymorphism, but, over an intercontinental scale, only non-delete trnL was found in Africa and the USA. ISSR analysis within each population detected a higher genetic distance between the samples with different trnL haplotypes, suggesting the presence of two different genetic lineages within this species, geographically overlapping in the Mediterranean Basin.     

Chloroplast phylogeographic patterns of Calligonum sect. Pterococcus (Polygonaceae) in arid Northwest China

To understand the impacts of past climatic change and geological events on the evolutionary history of Calligonum sect. Pterococcus, including C. aphyllum, C. rubicundum and C. leucocladum, a total of 128 individuals from 14 populations, mainly from arid Northwest China, were sampled. Two cpDNA intergenic spacer regions (rpl32‐trnL and ycf6‐psbM) were sequenced and 11 haplotypes were identified. Levels of genetic differentiation between populations was low in C. rubicundum (F_ST = 0.54317, p < 0.001) and C. aphyllum (F_ST = 0.55795), while much higher in C. leucocladum (F_ST = 0.95800, p < 0.001), possibly as an effct of differences in geographic distributions and habitats. Analysis of molecular variance (AMOVA) revealed that most of the total genetic variations occurred among species (72.97%). Among eleven identified haplotypes, only H1 and H2 were shared between C. aphyllum and C. rubicundum, while nine were private for one of the three species. The eleven identified haplotypes were divided into two major clades, but they did not yield three species‐specific lineages. Calligonum sect. Pterococcus therefore not appeared reciprocally monophyletic, more likely due to incomplete lineage sorting than hybridization. Mismatch distribution analysis suggested that only C. aphyllum has experienced recent demographic expansion. Divergence time among the 11 haplotypes was estimated at between 2.84 Ma and 0.06 Ma. Within the two clades, haplotype divergence began in early Pleistocene and mainly occurred during the middle to late Pleistocene and was most likely triggered by Quaternary climatic oscillations and increasing aridity of the region.     

Colonization and diversification of the spider genus Pholcus Walckenaer, 1805 (Araneae, Pholcidae) in the Macaronesian archipelagos: evidence for long-term occupancy yet rapid recent speciation

Macaronesian archipelagos stand apart from other oceanic islands reputed as laboratories for the study of evolution by their proximity to the mainland, lack of subsidence, and steep ecological gradients. The genus Pholcus Walckenaer, 1805, commonly known as daddy-long-leg spiders, is one of the most speciose arthropod groups in the region, with 25 endemic species. In the present study, we use information from four mitochondrial genes, along with morphological data, to examine the phylogenetic relationships and diversification patterns of the genus in the region. Phylogenetic analyses support monophyly of Macaronesian Pholcus including the Moroccan species Pholcus vachoni and hence a single colonization event in the archipelagos. Madeira colonizers most likely originated from the Canaries, and a back-colonization of the nearby mainland receives further support. Estimated lineage divergence times suggest a long-time presence of Pholcus in the region, but also reveal that most present-day species are the result of recent, and probably rapid, speciation events. Diagnostic characters among Macaronesian Pholcus are confined to structures involved in copulation. Coupled with the extremely high diversification rate, the highest recorded for spiders, these copulatory characters suggest that sexual selection has played a key role in the local diversification of Pholcus in Macaronesia.     

Genetic diversity and maternal origin of Northeast African and South American donkey populations

To investigate the mtDNA variation and origin of maternal lineages in South American donkeys and to reassess the domestication of donkeys in northeast Africa, we analyzed sequences (489 bp of the D‐loop) from 323 domestic donkeys sampled from Peru, Brazil, Ethiopia and Egypt. Altogether, the 323 sequences displayed 53 different haplotypes (45 in Ethiopia, 14 in Egypt, eight in Peru and six in Brazil). Among the four populations, Egyptian donkeys possessed the highest haplotype diversity (0.910 ± 0.032), followed by Brazilian donkeys (0.879 ± 0.060). The Clade I haplotypes dominated in Peruvian donkeys (65%), whereas Clade II haplotypes dominated in Brazilian donkeys (67%). Estimates of F_ST values showed a high genetic differentiation between Peruvian and Brazilian donkey populations (F_ST = 0.4066), which could be explained by the complex introduction history of South American donkeys. Phylogeographic analysis indicates that northeast Africa could be the most probable domestication center for Clade I donkeys. Analysis of molecular variance confirmed a weak genetic structure in domestic donkey populations among four continents (Europe, Asia, Africa and South America).     

Genetic divergence and evidence for sympatric host-races in the highly polyphagous brown tail moth,Euproctis chrysorrhoea (Lepidoptera: Erebidae)

The brown tail moth (BTM) Euproctis chrysorrhoea (Linnaeus 1758) (Lepidoptera: Erebidae) is a forest and ornamental pest in Europe and the United States. Its extreme polyphagy, and documented phenological shift associated with host use suggest the presence of distinct host-races. To test this hypothesis, we sampled BTM infesting different host species in several locations along its distribution, and used DNA sequence data (a total of 1,672 bp from cytochrome c oxidase subunit I, elongation factor 1-alpha, and wingless) to produce haplotype networks and reconstruct the phylogenetic relationships between individuals. Population genetic diversity indices pointed out a higher genetic diversity in Europe, particularly in the samples from southern Spain and southern England. Lower F _ST values were found between geographically closer populations when compared to more distant ones, but analyses of molecular variance and Mantel tests failed to reveal geographically associated genetic differentiation. However, haplotype networks and phylogenetic reconstructions revealed a previously unknown genetic differentiation within the BTM, with one lineage circumscribed to southern Europe. Although BTM haplotypes did not cluster according to their host plant, host-associated haplotypes were observed within certain geographic regions. Hence, our data support the existence of host-races of BTM within southern Spain and southern England, where populations from different hosts occur in sympatry.     

A reconstruction of Palaeo‐Macaronesia,with particular reference to the long‐term biogeography of the Atlantic island laurel forests

Macaronesia is a biogeographical region comprising five Atlantic Oceanic archipelagos: the Azores, Madeira, Selvagen (Savage Islands), Canaries and Cape Verde. It has strong affinities with the Atlantic coast of the Iberian Peninsula and the north‐western fringes of Africa. This paper re‐evaluates the biogeographical history and relationships of Macaronesia in the light of geological evidence, which suggests that large and high islands may have been continuously available in the region for very much longer than is indicated by the maximum surface area of the oldest current island (27 Ma) – possibly for as long as 60 million years. We review this literature, attempting a sequential reconstruction of Palaeo‐Macaronesia from 60 Ma to the present. We consider the implications of these geological dynamics for our understanding of the history of colonization of the present islands of Macaronesia. We also evaluate the role of these archipelagos as stepping stones and as both repositories of palaeo‐endemic forms and crucibles of neo‐endemic radiations of plant and animal groups. Our principal focus is on the laurel forest communities, long considered impoverished relicts of the Palaeotropical Tethyan flora. This account is therefore contextualized by reference to the long‐term climatic and biogeographical history of Southern Europe and North Africa and by consideration of the implications of changes in land–sea configuration, climate and ocean circulation for Macaronesian biogeography. We go on to provide a synthesis of the more recent history of Macaronesian forests, which has involved a process of impoverishment of the native elements of the biota that has accelerated since human conquest of the islands. We comment briefly on these processes and on the contemporary status and varied conservation opportunities and threats facing these forests across the Macaronesian biogeographical region.     

Mitochondrial DNA variation in Spanish populations of Ceratitis capitata (Wiedemann) (Tephritidae) and the colonization process

Abstract: Mitochondrial DNA variation in the Mediterranean fruit fly Ceratitis capitata (Wiedemann, Diptera: Tephritidae) was studied in three natural populations from southern, central and eastern Spain by means of restriction fragment length polymorphisms using 22 restriction endonucleases. Nine different haplotypes were found based upon the restriction patterns of the seven polymorphic endonucleases, providing a measure of discrimination between populations (N_ST = 0.2462, F_ST = 0.154). The observed distribution of haplotypes, corroborated by a parsimonious unrooted tree, suggests an ancient origin for haplotype VII, and a first step in the colonization of the Iberian Peninsula and subsequently the northern and eastern Mediterranean basin, through the Straits of Gibraltar. No relationship can be established between the colonization process for Europe and America.