A Bryophyte Flora of Kent. IV. Addenda and Corrigenda

Abstract

Based on a revision of Andreaea from the Canary Islands, the occurence of two species is confirmed. A key is provided. A. crassifolia Luisier occurs on Gran Canaria, La Palma and Tenerife. A. heinemannii Hampe &; Müll.Hal. occurs on La Palma. Former records of A. heinemannii s.l. for Gran Canaria and Tenerife refer to A. crassifolia.     

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.     

Index to Vegetatio,volumes 74–110 (1988–1994)

Chamaecytisus proliferus (L.fil.) Link (Fabaceae: Genisteae) represents a species complex in the Canary Islands. Floristic data from 147 releves from the whole complex were collected and analysed by classification (TWINSPAN) and ordination (DECORANA) methods. Results indicate that white escobon of Tenerife, escobon of El Hierro, white escobon of Gran Canaria and typical tagasaste in La Palma are associated with those plant communities from the north of these islands which are under the influence of the north-eastern trade winds. Narrow-leaved escobon in Tenerife and La Gomera, escobon of southern Gran Canaria and white tagasaste of La Palma are found in those areas which are not under the direct influence of these winds. Morphological forms from the more easterly islands (Gran Canaria and Tenerife-La Gomera) have the broadest ecological range and they have played an important role in the floristic changes which have taken place after the destruction of the forests in these islands. The highest priorities for in situ conservation should be given to wild populations of typical tagasaste, white escobon of Tenerife and escobon of El Hierro.Abbreviations IBPGR International Board for Plant Genetic Resources - DECORANA Detrended Correspondence Analysis - OTU Operational Taxonomic Unit - TWINSPAN Two Way Indicator Species Analysis     

Delimiting species boundaries for endangered Canary Island grasshoppers based on DNA sequence data

Mitochondrial DNA cytochrome oxidase subunit I and nuclear ITS2 sequences were surveyed from Canary Islands threatened species of the genera Purpuraria and Acrostira (Orthoptera: Pamphagidae). Phylogenetic and population analyses show that the two previously recognized Purpuraria erna subspecies are not valid as conservation units, and that there is a new unrecognized species of Purpuraria, coincident with recently discovered morphological variation within the genus. In addition, mitochondrial introgression seems to occur between the two Purpuraria species in southwest Lanzarote. Species-delimitation based on the morphological taxonomy of Acrostira, which recognizes four single-island endemics, is only partially supported by the genetic data. It shows that currently admitted species from the central and western islands of Tenerife, La Gomera and La Palma are closely related, with evidence of recent gene flow between the Tenerife and La Gomera populations. MtDNA variation also showed that A. euphorbiae, currently considered as the most critically endangered grasshopper species in the Canaries, has lower population diversity than its close relatives.     

Predation of insects by feral cats (<Emphasis Type="Italic">Felis silvestris catus</Emphasis> L., 1758) on an oceanic island (La Palma,Canary Island)

Predation of insects by feral cats (Felis silvestris catus) on a heterogeneous oceanic island (La Palma, Canary Islands) was studied. A total of 127 invertebrates were identified in the analysis of 500 scats (100 from each habitat of the Island). Invertebrates appear in 18.00% of the scats, representing an insignificant percentage of the total consumed biomass by feral cats on La Palma Island (0.05%). Insects were the most common invertebrate prey both in percentage of occurrence (90.6%) and invertebrate biomass (93.53%), with a total of 115 prey items. Orthoptera, Lepidoptera and Coleoptera were the main prey groups. Among the five main habitats present in La Palma Island, the temperate forest shows the lowest consumption of invertebrates, although insect consumption did not show statistical differences. However Orthoptera and Lepidoptera were more frequently preyed on in the pine forest and in the xerophytic shrub, respectively. Moreover, applying the Simplified Morisita index, a different insect composition of the diet was observed among habitats. Although, none of insects predated by feral cats are threatened, the identification of invertebrate component of the feral cats’ diet is an important tool for the correct understanding of predation significance and to prevent damage to endangered insect species.     

Characterization of ten polymorphic microsatellite loci for the threatened species Grimmia curviseta Bouman (Grimmiaceae,Musci)

Grimmia curviseta Bouman is a moss species endemic to the Canary Islands in the North Atlantic Ocean, which has a very restricted distribution across the summit areas of Tenerife and La Palma islands. Using massive sequencing, we developed ten polymorphic microsatellite markers for this species. The pattern of microsatellite alleles per locus provides preliminary evidence that G. curviseta is allodiploid. The average number of alleles per locus ranged from 2 to 7, and observed heterozygosities varied from 0.022 to 1.000. A significant genetic differentiation was observed between the Tenerife and La Palma populations, suggesting that there are some limitations to dispersal. This set of microsatellites constitutes a valuable tool to investigate patterns of genetic diversity across the distribution range of G. curviseta, information that may eventually be used to establish conservation strategies for the species. Owing to the cross-amplification with the closely related species Grimmia montana Bruch & Schimp., our study also demonstrates the utility of these markers for population-level genetic analyses in the genus Grimmia.     

The evolutionary history of Afrocanarian blue tits inferred from genomewide SNPs

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

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.     

Microsatellite loci isolation in the Canarian common chaffinch (Fringilla coelebs) and their utility in other Canarian finches

The taxonomic classification of the common chaffinch (Fringilla coelebs) that inhabits the Canary Islands has been under debate for decades, mainly due to the absence of nuclear DNA analyses. In this study we describe the isolation and characterization of ten microsatellite loci (AAAG, AAAT and GT) from a La Palma specimen using an enrichment protocol. Two loci were monomorphic in the populations analysed (La Gomera and La Palma), but the remaining ones presented 2 or more alleles, with an average of 11.63 alleles per locus and an average observed heterozygosity of 0.735 (n = 44). All loci were tested for their utility in other Canarian populations and other finch species.     

Niche Occupation by Invasive Ground-Dwelling Predator Species in Canarian Laurel Forests

The distribution and co-occurrence of three groups of carnivorous soil macro-invertebrates (Carabidae, Staphylinidae, Chilopoda) were examined in laurel forests of the western Canary Islands. The species numbers per site decreased from East to West (La Gomera, El Hierro, La Palma) in both, beetles and centipedes. No evidence was found for the ‘diversity-invasibility-hypothesis’ sensu Elton. The number of invasive species per site increased with that of native species in Chilopoda, and was not significant in Carabidae+Staphylinidae. Carabidae and Staphylinidae were combined to form a guild of non-specialized ground-dwelling predatory beetles. The mandible length of adults and larvae was used as an indicator of the preferred food size class to determine the food niche width and niche separation. Two invasive coleopteran species were also examined: Ocypus olens occupied the vacant top predator niche on El Hierro, and Laemostenus complanatus occupied the vacant medium size predator niche on La Palma. Neither of these species was found in laurel forests of any other island where these niches are occupied by autochthonous species, though they are introduced on these islands too. The Chilopoda occurred in the forests with seven invasive and seven native species. Autochthonous and introduced centipedes species of the same size class and group are mutually exclusive.     

Drosophilid assemblages as a bioindicator system of human disturbance in the Brazilian Savanna

According to the Convention on Biological Diversity (CBD) the development of bioindicators is extremely necessary to achieve the conservation targets by 2010, and insects are considered an effective group for this goal. Drosophilids are regarded as potential indicators, although this idea remains untested. Therefore, we followed up a protocol to test the drosophilid potential indicator for human disturbance in the Brazilian Savanna, one of the richest and most threatened tropical biomes in the world. Sampling was undertaken in one urban environment and two biological reserves, representing four habitat types (undisturbed gallery forest, disturbed gallery forest, undisturbed savanna, and urban environment). We examined differences in the drosophilid assemblages among habitat types and used the Indicator Value (IndVal) method to point out the indicator species. We also tested the two-stage indicator validation, a protocol recently proposed in the literature, to validate the indicator species for undisturbed gallery forest and savannas, in independent samples. The assemblage variables varied mainly in undisturbed gallery forests, and reflected changes from an undisturbed to a disturbed stage. The IndVal associated with the two-stage protocol showed reliable characteristic species, which are very helpful for diagnostic surveys. Likewise, species that can detect changes in the habitats were also found. We found a set of indicators, which together may be very efficient for both assessing and reflecting a variety of conditions, improving the confidence of the bioindication system, expanding the taxonomic options for bioindicators, and therefore, contributing to the conservation of this region.     

Batch fecundity of Lutjanus carponotatus (Lutjanidae) and implications of no-take marine reserves on the Great Barrier Reef, Australia

This study investigated body size to fecundity relationships of a reef fish species targeted by line fishing, and examines the potential benefits of increased batch fecundity in no-take reserves compared to fished areas around the Palm, Whitsunday and Keppel Island Groups, Great Barrier Reef, Australia. Lutjanus carponotatus batch fecundity increased with fork length in a non-linear relationship that was best described by a power function. Batch fecundity differed by more than 100-fold among individuals, with a range from 7,074 to 748,957 eggs in fish ranging from 184 to 305 mm fork length. Furthermore, egg diameter increased with fish size. Based on underwater visual census, the potential batch fecundity per unit area in all three island groups ranged from 1.0 to 4.2 times greater in the no-take reserves than in the fished areas between 2001 and 2004. In 2002, a mean 2.3-fold difference in biomass between no-take reserves and fished areas converted to a mean 2.5-fold difference in batch fecundity per unit area. Greater batch fecundity, longer spawning seasons and potentially greater larval survival due to larger egg size from bigger individuals might significantly enhance the potential benefits of no-take marine reserves on the Great Barrier Reef.     

Cactophilic Drosophila in South America: A Model for Evolutionary Studies

The Drosophila buzzatii cluster is composed of seven cactophilic species and their known geographical distribution encompasses the open vegetation diagonal, which includes the morphoclimatic Domains of the Caatinga, Chaco and Cerrado, which are situated between the Amazon and the Atlantic forests. Besides these areas, these cactophilic species are also found in a narrow strip along the Atlantic coast from northeastern Brazil to the southern tip of the country. The hypothesis of vicariant events, defining the core areas of each species, is proposed to explain the historical diversification for the cluster. The intraspecific analysis for the cluster shows a population structure with gene flow restricted by distance, range expansion with secondary contact resulting in introgression and simpatry, especially in the limits of the species distribution, polytypic populations and assortative mating in inter population experiments. There is a variation related to these events that depends on the species and geographic origin of the population analyzed. These events are, hypothetically, described as the results of expansion and retraction of the population ranges, as a consequence of their association with cacti, which theoretically follow the expansion and retraction of dry areas during the paleoclimatic oscillations in South America, as that promoted by the glacial cycles of the Quaternary. The Drosophila buzzatii cluster is divided into two groups. The first one is composed of D. buzzatii, a species that has a broad geographic distribution and no significant differentiation between its populations. The second is the Drosophila serido sibling set, which encompasses the others species and is characterized by a significant potential for differentiation.     

Reconciling gene trees with organism history: the mtDNA phylogeography of three Nesotes species (Coleoptera: Tenebrionidae) on the western Canary Islands

The processes of island colonization and speciation are investigated through mtDNA studies on Canary Island beetles. The genus Nesotes (Coleoptera: Tenebrionidae) is represented by 19 endemic species on the Canary Islands, the majority of which are single island endemics. Nesotes conformis is the most widespread, occurring on Gran Canaria, Tenerife, La Palma and El Hierro. Nesotes conformis forms a paraphyletic assemblage, with a split between Gran Canaria and the other three islands. Nesotes conformis of the western Canary Islands cluster with Nesotes altivagans and Nesotes elliptipennis from Tenerife. Fifty‐two individuals from this western islands species complex have been sequenced for 675 base pairs of the mtDNA cytochrome oxidase II gene, representing Tenerife, La Palma and El Hierro. A neighbour joining analysis of maximum likelihood distances resulted in three distinct mtDNA lineages for N. conformis, two of which also include mitotypes of N. altivagans and N. elliptipennis. Through application of parametric bootstrap tests, we are able to reject hypotheses of monophyly for both N. conformis and N. altivagans. Nesotes altivagans and N. elliptipennis are poorly separated morphologically and mtDNA sequence data adds support to this being one species with a highly variable morphology. We propose that N. altivagans/N. elliptipennis is recently derived from two ancestral mtDNA lineages within N. conformis from the Teno region of Tenerife. We further propose colonization of the younger islands of La Palma and El Hierro by N. conformis from a mitochondrial lineage within the Teno massif (colonization; diversification; mitochondrial DNA; Canary Islands; Coleoptera).     

Mitochondrial DNA phylogeography and population history of Meladema diving beetles on the Atlantic Islands and in the Mediterranean basin (Coleoptera,Dytiscidae)

The phylogeny and population history of Meladema diving beetles (Coleoptera, Dytiscidae) were examined using mitochondrial DNA sequence from 16S ribosomal RNA and cytochrome oxidase I genes in 51 individuals from 22 populations of the three extant species (M. imbricata endemic to the western Canary Islands, M. lanio endemic to Madeira and M. coriacea widespread in the Western Mediterranean and on the western Canaries), using a combination of phylogenetic and nested clade analyses. Four main lineages are observed within Meladema, representing the three recognized species plus Corsican populations of M. coriacea. Phylogenetic analyses demonstrate the sister relationship of the two Atlantic Island taxa, and suggest the possible paraphyly of M. coriacea. A molecular clock approach reveals that speciation within the genus occurred in the Early Pleistocene, indicating that the Atlantic Island endemics are not Tertiary relict taxa as had been proposed previously. Our results point to past population bottlenecks in all four lineages, with recent (Late-Middle Pleistocene) range expansion in non-Corsican M. coriacea and M. imbricata. Within the Canary Islands, M. imbricata seems to have independently colonized La Gomera and La Palma from Tenerife (although a colonization of La Palma from La Gomera cannot be discarded), and M. coriacea has independently colonized Tenerife and Gran Canaria from separate mainland lineages. In the Mediterranean basin this species apparently colonized Corsica on a single occasion, relatively early in its evolutionary history (Early Pleistocene), and has colonized Mallorca recently on multiple occasions. On the only island where M. coriacea and M. imbricata are broadly sympatric (Tenerife), we report evidence of bidirectional hybridization between the two species.     

Large Bird Declines with Increasing Human Pressure in Savanna Woodlands (Burkina Faso)

The impacts of human pressure, habitat changes, and efficiency of conservation measures on the non-passerine bird community were assessed in the Sudanian belt of Burkina Faso. Three regions with increasingly large protected areas and higher conservation status were compared. In each region, natural savanna woodlands were also compared to traditionally cultivated areas. Six groups of diurnal raptors and six groups of other large bird species were counted along 167 5-km transect counts during the dry season. Within savanna woodlands, the only significant and consistent declines from the little disturbed Eastern National Parks to the more populated and exploited Western forest reserves were those of the largest terrestrial game birds (Guinea fowl, Bustard, Ground Hornbill), eagles and vultures, which became virtually extinct in the least protected areas. Although some species groups maintained substantial, if not higher populations in cultivated areas (rollers, non-game terrestrial birds, several raptors), the overall abundance of both raptor and non-raptor species declined markedly (50–65%) from natural savanna woodlands to tree-dotted cultivated and fallow fields. Again large game birds, eagles and vultures were absent from every agricultural areas censused. Yet, all of them were still commonly recorded in the same natural and cultivated areas during surveys I conducted in the same regions from 1968 to 1973. The main factors correlated with large bird extinctions or declines from national parks to mere forest reserves and from natural woodlands to fields were first related to hunting, then to habitat degradation and fragmentation through intensive cattle grazing and wood cutting and extension of cultivated areas.     

Pollination of Habenaria tridactylites on the Canary Islands

We investigated the pollination of Habenaria tridactylites, an endemic orchid of the Canary Islands. The entirely green, widely open flowers have a long spur containing nectar. We carried out fieldwork, a molecular clock analysis, herbarium surveys, identified pollinators by both morphology and DNA barcoding, and measured the length of floral spurs and insect tongues using a combination of traditional and innovative micro‐CT scanning methods to 1) determine the pollinator of this orchid and 2) investigate correlations between local mean spur length and age, altitude and longitude of the island. Habenaria tridactylites was found to be pollinated on Tenerife by both small and intermediate sized moth species with variable tongue lengths and mostly belonging to Geometridae and to a lesser extent Crambidae, Erebidae, Noctuidae and Tortricidae. Of the sixteen moth species identified, nine are endemic to the Canary Islands or Macaronesia. The different local populations of H. tridactylites on the islands of Gran Canaria, El Hierro, La Gomera, La Palma and Tenerife with different ages and distances from mainland Africa, did not show a significant correlation of mean spur length and altitude, but did show a significant and positive linear correlation with longitude and the geological age of the island. The latter is congruent with the evolutionary arms race theory first proposed by Darwin, suggesting that flowers gradually evolve longer spurs and pollinators longer tongues.     

Ecological niche modelling confirms potential north‐east range expansion of the nine‐banded armadillo (Dasypus novemcinctus) in the USA

The distribution of the nine banded armadillo (Dasypus novemcinctus), the only species in the family Dasypodidae found in the USA, has expanded greatly since the species was first recorded in southern Texas in 1849. Currently, the range of D. novemcinctus includes 15 states in the USA. Previous studies on the geographical expansion of this species, based on physiological experiments and distribution surveys, revealed a possible western moisture limit, a northern temperature limit, and potential north‐eastward range expansion in the USA. We applied an ecological niche modelling approach and produced a potential distribution map of D. novemcinctus with comparable western (102 °W) and northern (40 °N) limits, and confirmed the possibility of further north‐east range expansion to climatically suitable areas in the USA.     

Numerical analyses of RAPD data highlight the origin of cultivated tagasaste (Chamaecytisus proliferus ssp. palmensis) in the Canary Islands

Random Amplified Polymorphic DNA (RAPD) was used to generate molecular markers to trace the origin of the fodder legume tagasaste (Chamaecytisus proliferus (L. fil.) Link ssp. palmensis (H. Christ) Kunkel) in the Canary Islands. Results from multivariate analyses of data through Two Way Indicator Species Analysis (TWINSPAN) and Detrended Correspondence Analysis (DECORANA) showed that genotypes collected on the island of La Palma exhibited a wider range of variation than those from the other islands. This supports the existing hypothesis that tagasaste originated on La Palma and emphasizes the importance of conserving and evaluating germ plasm from this island.     

Modelling the potential distribution of the invasive tomato red spider mite, <Emphasis Type="Italic">Tetranychus evansi</Emphasis> (Acari: Tetranychidae)

Predicting the potential geographical distribution of a species is particularly important for pests with strong invasive abilities. Tetranychus evansi Baker & Pritchard, possibly native to South America, is a spider mite pest of solanaceous crops. This mite is considered an invasive species in Africa and Europe. A CLIMEX model was developed to predict its global distribution. The model results fitted the known records of T. evansi except for some records in dry locations. Dryness as well as excess moisture stresses play important roles in limiting the spread of the mite in the tropics. In North America and Eurasia its potential distribution appears to be essentially limited by cold stress. Detailed potential distribution maps are provided for T. evansi in the Mediterranean Basin and in Japan. These two regions correspond to climatic borders for the species. Mite establishment in these areas can be explained by their relatively mild winters. The Mediterranean region is also the main area where tomato is grown in open fields in Europe and where the pest represents a threat. According to the model, the whole Mediterranean region has the potential to be extensively colonized by the mite. Wide expansion of the mite to new areas in Africa is also predicted. Agricultural issues highlighted by the modelled distribution of the pest are discussed. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.