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

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

Genetically depauperate in the continent but rich in oceanic islands: Cistus monspeliensis (Cistaceae) in the Canary Islands

Background

Population genetic theory holds that oceanic island populations are expected to have lower levels of genetic variation than their mainland counterparts, due to founder effect after island colonization from the continent. Cistus monspeliensis (Cistaceae) is distributed in both the Canary Islands and the Mediterranean region. Numerous phylogenetic results obtained in the last years allow performing further phylogeographic analyses in Cistus.

Methodology/Principal Findings

We analyzed sequences from multiple plastid DNA regions in 47 populations of Cistus monspeliensis from the Canary Islands (21 populations) and the Mediterranean basin (26 populations). The time-calibrated phylogeny and phylogeographic analyses yielded the following results: (1) a single, ancestral haplotype is distributed across the Mediterranean, whereas 10 haplotypes in the Canary Islands; (2) four haplotype lineages are present in the Canarian Islands; (3) multiple colonization events across the archipelago are inferred; (4) the earliest split of intraspecific lineages occurred in the Early to Middle Pleistocene (<930,000 years BP).

Conclusions/Significance

The contrasting pattern of cpDNA variation is best explained by genetic bottlenecks in the Mediterranean during Quaternary glaciations, while the Canarian archipelago acted as a refugium of high levels of genetic diversity. Active colonization across the Canarian islands is supported not only by the distribution of C. monspeliensis in five of the seven islands, but also by our phylogeographic reconstruction in which unrelated haplotypes are present on the same island. Widespread distribution of thermophilous habitats on every island, as those found throughout the Mediterranean, has likely been responsible for the successful colonization of C. monspeliensis, despite the absence of a long-distance dispersal mechanism. This is the first example of a plant species with higher genetic variation among oceanic island populations than among those of the continent.     

Historical Isolation of the Galápagos Carpenter Bee (Xylocopa darwini) despite Strong Flight Capability and Ecological Amplitude

Colonization across the Galápagos Islands by the carpenter bee (Xylocopa darwini) was reconstructed based on distribution of mitochondrial haplotypes (cytochrome oxidase II (COII) sequences) and haplotype lineages. A total of 12 haplotypes were found in 118 individuals of X. darwini. Distributional, phylogenetic and phylogeographic analyses suggest early colonization of most islands followed by historical isolation in two main groups: eastern and central-western islands. Evidence of recurrent inter-island colonization of haplotypes is largely lacking, despite strong flight capability and ecological amplitude of the species. Recent palaeogeographic data suggest that several of the current islands were connected in the past and thus the isolation pattern may have been even more pronounced. A contrast analysis was also carried out on 10 animal groups of the Galápagos Islands, and on haplotype colonization of seven animal and plant species from several oceanic archipelagos (the Galápagos, Azores, Canary Islands). New colonization metrics on the number of potential vs. inferred colonization events revealed that the Galápagos carpenter bee shows one of the most significant examples of geographic isolation.     

Inferring native range and invasion scenarios with mitochondrial DNA: the case of T. solanivora successive north�Csouth step-wise introductions across Central and South America

Tecia solanivora (Lepidoptera: Gelechiidae) is an invasive species that attacks the tubers of the potato Solanum tuberosum. It is considered a serious pest of potato crops and stocks in all countries where it occurs. In the present study, we sequenced 541 individuals sampled across the T. solanivora distribution range, using the mitochondrial DNA marker Cytochrome b (Cytb) to delimit the area of species origin. We also analyzed the genetic structure of T. solanivora in its putative area of origin and described differences in haplotype diversity between samples from different geographic origins affected by the invasion. We observed a gap in the level of genetic diversity between Guatemalan samples (h between 0.77 and 0.97) and those from Costa-Rica, Venezuela, Colombia, Ecuador and the Canary Islands (h between 0 and 0.56). The number of haplotypes has decreased over the colonization process, ending with the observation of a single haplotype in Colombia, Ecuador and the Canary Islands. Consequently, the invasion of South American countries by T. solanivora is likely to have had a front-like step-wise progression, where the most recently invaded country becomes the source of subsequent invasion.     

Trophic ecology of feral cats (Felis silvestris f. catus) in the main environments of an oceanic archipelago (Canary Islands): An updated approach

The diet of feral cats in the main habitats of the Canary Islands is composed of introduced mammals, birds, reptiles and insects. However, introduced mammals constitute the main source of biomass consumed, followed in importance by reptiles and birds. PCA analysis of biomass revealed the ordination of three different groups, corresponding to the diet in the laurel forest (La Gomera), thermophilous forest (El Hierro) and one large group that include the rest of habitat types. A similar pattern was observed when these habitats were analyzed in a single island (La Palma). Oryctolagus cuniculus was an important prey in practically all habitats, while Rattus rattus was frequently captured in the laurel forest, Mus musculus domesticus in the open shrubs (both xeric and high mountain), reptiles (mainly lizards genus Gallotia) in the open habitats of Tenerife, birds play a relative role in forest habitats, and large invertebrates (basically Orthoptera and Coleoptera) in the three forest habitats and in the xerophytic shrub of Fuerteventura. Morisita's index of similarity of diet showed maximum differences between the forest habitats (pine and thermophilous vs. laurel forest), indicating an important heterogeneity in the diet of feral cats in these environments. Shrub habitats showed smaller values of Levin's niche breadth than those from the forest habitats, showing a broader diet in the latter. Lastly, the diet of feral cats on the Canary Islands follows the general pattern of other islands located at similar latitude and mainly composed by rabbits and mice. However, specific preys such as lizards, rats or birds, play an important role in particular habitats in which they are abundant.     

Mitochondrial DNA variability in the Canary Islands honeybees (Apis mellifera L.)

The mitochondrial DNA (mtDNA) of individuals from 79 colonies of Apis mellifera from five Canary Islands was studied using the Dra I test based on the restriction of PCR products of the tRNAleu–COII intergenic region. Five haplotypes of the African (A) lineage and one of the west European (C) lineage were found. The haplotypes A14 and A15 are described for the first time. These haplotypes have a new P sequence named P₁. The wide distribution and high frequency of haplotype A15 suggest that it is characteristic of the Canarian Archipelago. Sources of haplotype variability of honeybee mtDNA in the Canary Islands (waves of colonization from Africa, queen importations, habitat diversification) are discussed.     

Strains nodulating Lupinus albus on different continents belong to several new chromosomal and symbiotic lineages within Bradyrhizobium

In this work we analysed different chromosomal and symbiotic markers in rhizobial strains nodulating Lupinus albus (white lupin) in several continents. Collectively the analysis of their rrs and atpD genes, and 16S-23S intergenic spacers (ITS), showed that they belong to at least four chromosomal lineages within the genus Bradyrhizobium. Most isolates from the Canary Islands (near to the African continent) grouped with some strains isolated on mainland Spain and were identified as Bradyrhizobium canariense. These strains are divided into two ITS subgroups coincident with those previously described from isolates nodulating Ornithopus. The remaining strains isolated on mainland Spain grouped with most isolates from Chile (American continent) forming a new lineage related to Bradyrhizobium japonicum. The strains BLUT2 and ISLU207 isolated from the Canary Islands and Chile, respectively, formed two new lineages phylogenetically close to different species of Bradyrhizobium depending on the marker analyzed. The analysis of the nodC gene showed that all strains nodulating L. albus belong to the biovar genistearum; nevertheless they form four different nodC lineages of which lineage C is at present exclusively formed by L. albus endosymbionts isolated from different continents.     

Invasion by Rattus rattus into native coastal forests of south‐eastern Australia: are native small mammals at risk?

The black rat, Rattus rattus, is an alien rodent in Australian ecosystems where niche overlap with native small mammals may lead to competition for resources and displacement of native species. In coastal habitats surrounding Jervis Bay in south‐eastern Australia, R. rattus co‐occurs with the native bush rat, Rattus fuscipes, and brown antechinus, Antechinus stuartii. Relative distributions and abundances, and fine‐scale space use suggest invasive and native rodents compete for use of space and habitat. Such competitive interactions were not evident between R. rattus and native A. stuartii, which was negatively influenced more by disturbance to habitat. Differences in rodent communities between spatially separate forests forming the northern and southern peninsulas of Jervis Bay potentially reflect symmetrical competition and differences in competitive outcomes. In southern forests, R. rattus was largely restricted to patches of disturbed forest associated with campgrounds. Competitive interference by native rodent populations inhabiting surrounding intact forests may have so far limited R. rattus colonization of these areas. In northern forests, R. rattus was the predominant rodent irrespective of disturbance, while populations of R. fuscipes were unusually low seemingly due to poor juvenile recruitment. Native individuals avoided areas frequented by adult R. rattus and given that species did not partition use of microhabitats, R. rattus most likely precluded R. fuscipes from suitable habitat and in doing so limited native populations. We discuss how natural disturbance of habitat and human activity have potentially facilitated successful invasion by R. rattus of the northern forests. Studies that manipulate rodent populations are required to support these interpretations of observed patterns.     

Focus: Zoonotic Disease: Prevalence and Diversity of the Streptobacillus Rat-bite Fever Agent,in Three Invasive,Commensal Rattus Species from South Africa

Rat-bite fever is an over-looked, global zoonotic disease that has a mortality rate of up to 13%, if untreated. Historically, this rat-borne disease has been attributed to one of two causative agents, Streptobacillus moniliformis or Spirillum minus. Given the confirmed presence of multiple invasive Rattus host species, high rat densities in urban, informal human settlements and increasing reports of rat bites in South Africa, we undertook a retrospective assessment of Streptobacillus in rats sampled from 16 urban sites, in Gauteng, the smallest but most populous Province in South Africa. Using a multi-gene PCR-sequencing approach, we confirmed Streptobacillus presence in 50.9% of oral swabs from three rat species and the presence of two Streptobacillus species, viz. S. moniliformis and S. notomytis. The two members of the cryptic Rattus rattus species complex (R. rattus and R. tanezumi), which are morphologically indistinguishable from each other, had markedly different colonization rates. Whereas 48.6% of rats from this species complex were Streptobacillus-positive, only 32.3% of Rattus tanezumi were positive compared to 61.5% R. rattus. Rattus norvegicus had an intermediate prevalence of 55.6%. Phylogenetic analysis of four gene regions (16S rRNA, gyrB, groEL, recA) identified two discrete lineages; S. moniliformis occurred exclusively in R. norvegicus, and S. notomytis was restricted to the two members of the R. rattus species complex; this represents the first report of Streptobacillus in R. tanezumi. These results highlight a largely overlooked zoonotic threat posed by invasive rats and confirm the presence of two discrete and potentially host-specific Streptobacillus lineages in South Africa.     

Invasion facilitates hybridization with introgression in the Rattus rattus species complex

Biological invasions result in novel species interactions, which can have significant evolutionary impacts on both native and invading taxa. One evolutionary concern with invasions is hybridization among lineages that were previously isolated, but make secondary contact in their invaded range(s). Black rats, consisting of several morphologically very similar but genetically distinct taxa that collectively have invaded six continents, are arguably the most successful mammalian invaders on the planet. We used mitochondrial cytochrome b sequences, two nuclear gene sequences (Atp5a1 and DHFR) and nine microsatellite loci to examine the distribution of three invasive black rat lineages (Rattus tanezumi, Rattus rattus I and R. rattus IV) in the United States and Asia and to determine the extent of hybridization among these taxa. Our analyses revealed two mitochondrial lineages that have spread to multiple continents, including a previously undiscovered population of R. tanezumi in the south‐eastern United States, whereas the third lineage (R. rattus IV) appears to be confined to Southeast Asia. Analyses of nuclear DNA (both sequences and microsatellites) suggested significant hybridization is occurring among R. tanezumi and R. rattus I in the United States and also suggest hybridization between R. tanezumi and R. rattus IV in Asia, although further sampling of the latter species pair in Asia is required. Furthermore, microsatellite analyses suggest unidirectional introgression from both R. rattus I and R. rattus IV into R. tanezumi. Within the United States, introgression appears to be occurring to such a pronounced extent that we were unable to detect any nuclear genetic signal for R. tanezumi, and a similar pattern was detected in Asia.     

GEOTROPIC ORIENTATION OF YOUNG RATS

The geotropic orientation of Rattus rattus (roof rat) obeys the equations previously found applicable for Rattus norvegicus. The former is more sensitive, geotiopically, and the numerical values of the constants in the equations for the two forms are found to differ significantly. Certain consequences of this difference are pointed out.     

The origin of Rattus rattus on the Îles Éparses,Western Indian Ocean

Tollenaere et al. (Journal of Biogeography, 2010, 37 , 398–410) present a phylogeographic analysis of Rattus rattus for the Western Indian Ocean, with particular emphasis on Madagascar, but do not include samples from three island groups centrally located in the Mozambique Channel. Haplotypes from these islands provide additional information on the colonization pathways of R. rattus in the Western Indian Ocean region. For each of the three Îles Éparses groups in the Mozambique Channel, we test the competing hypotheses that colonization by R. rattus was most likely: (1) from the Arabian Peninsula, (2) from East Africa, (3) from Madagascar, or (4) from independent shipping. These results are combined with historical observations of the presence of R. rattus on these islands to give stronger inference on the colonization pathways. Additionally, more accurate colonization dates provide guidance for contemporary conservation management.     

Genetic variation in the kakerori (Pomarea dimidiata), an endangered endemic bird successfully recovering in the Cook Islands

The Cook Islands endemic kakerori (Pomarea dimidiata) underwent a severe population decline following the introduction of ship rats (Rattus rattus) in the late 1800s. By 1989, the sole population on Rarotonga consisted of 29 known birds. Subsequent intensive management efforts enabled this population to recover to around 250?C300 birds in recent years. This study, using microsatellite and mitochondrial DNA markers, assesses the level of genetic diversity and the genetic structure of the contemporary kakerori population on Rarotonga. No mitochondrial control region and cytochrome b haplotype diversity was found in the 11 samples examined at each locus. In 81 samples genotyped at 7 polymorphic microsatellite loci, an average of 4 alleles per locus were found, with an average observed heterozygosity of 0.65. No subpopulation division was found in this population. There was no evidence of inbreeding, but genetic bottleneck tests showed that the population had indeed experienced a significant genetic bottleneck. Recovery of the kakerori was successful in the past two decades despite low genetic diversity in terms of allelic diversity. Our data suggested that low allelic diversity did not hamper population expansion and the continued survival of this species, however, longer-term effects are still possible.     

Phylogeography of the introduced species Rattus rattus in the western Indian Ocean,with special emphasis on the colonization history of Madagascar

Aim To describe the phylogeographic patterns of the black rat, Rattus rattus, from islands in the western Indian Ocean where the species has been introduced (Madagascar and the neighbouring islands of Réunion, Mayotte and Grande Comore), in comparison with the postulated source area (India). Location Western Indian Ocean: India, Arabian Peninsula, East Africa and the islands of Madagascar, Réunion, Grande Comore and Mayotte. Methods Mitochondrial DNA (cytochrome b, tRNA and D‐loop, 1762 bp) was sequenced for 71 individuals from 11 countries in the western Indian Ocean. A partial D‐loop (419 bp) was also sequenced for eight populations from Madagascar (97 individuals), which were analysed in addition to six previously published populations from southern Madagascar. Results Haplotypes from India and the Arabian Peninsula occupied a basal position in the phylogenetic tree, whereas those from islands were distributed in different monophyletic clusters: Madagascar grouped with Mayotte, while Réunion and Grand Comore were present in two other separate groups. The only exception was one individual from Madagascar (out of 190) carrying a haplotype that clustered with those from Réunion and South Africa. ‘Isolation with migration’ simulations favoured a model with no recurrent migration between Oman and Madagascar. Mismatch distribution analyses dated the expansion of Malagasy populations on a time‐scale compatible with human colonization history. Higher haplotype diversity and older expansion times were found on the east coast of Madagascar compared with the central highlands. Main conclusions Phylogeographic patterns supported the hypothesis of human‐mediated colonization of R. rattus from source populations in either the native area (India) or anciently colonized regions (the Arabian Peninsula) to islands of the western Indian Ocean. Despite their proximity, each island has a distinct colonization history. Independent colonization events may have occurred simultaneously in Madagascar and Grande Comore, whereas Mayotte would have been colonized from Madagascar. Réunion was colonized independently, presumably from Europe. Malagasy populations may have originated from a single successful colonization event, followed by rapid expansion, first in coastal zones and then in the central highlands. The congruence of the observed phylogeographic pattern with human colonization events and pathways supports the potential relevance of the black rat in tracing human history.     

Demographic history of Canary Islands male gene-pool: replacement of native lineages by European

Background  

The origin and prevalence of the prehispanic settlers of the Canary Islands has attracted great multidisciplinary interest. However, direct ancient DNA genetic studies on indigenous and historical 17^th–18^th century remains, using mitochondrial DNA as a female marker, have only recently been possible. In the present work, the analysis of Y-chromosome polymorphisms in the same samples, has shed light on the way the European colonization affected male and female Canary Island indigenous genetic pools, from the conquest to present-day times.     

Scrophularia arguta,a widespread annual plant in the Canary Islands: a single recent colonization event or a more complex phylogeographic pattern?

Many studies have addressed evolution and phylogeography of plant taxa in oceanic islands, but have primarily focused on endemics because of the assumption that in widespread taxa the absence of morphological differentiation between island and mainland populations is due to recent colonization. In this paper, we studied the phylogeography of Scrophularia arguta, a widespread annual species, in an attempt to determine the number and spatiotemporal origins of dispersal events to Canary Islands. Four different regions, ITS and ETS from nDNA and psbA‐trnH and psbJ‐petA from cpDNA, were used to date divergence events within S. arguta lineages and determine the phylogenetic relationships among populations. A haplotype network was obtained to elucidate the phylogenetic relationships among haplotypes. Our results support an ancient origin of S. arguta (Miocene) with expansion and genetic differentiation in the Pliocene coinciding with the aridification of northern Africa and the formation of the Mediterranean climate. Indeed, results indicate for Canary Islands three different events of colonization, including two ancient events that probably happened in the Pliocene and have originated the genetically most divergent populations into this species and, interestingly, a recent third event of colonization of Gran Canaria from mainland instead from the closest islands (Tenerife or Fuerteventura). In spite of the great genetic divergence among populations, it has not implied any morphological variation. Our work highlights the importance of nonendemic species to the genetic richness and conservation of island flora and the significance of the island populations of widespread taxa in the global biodiversity.     

Cryptic diversity in Black rats Rattus rattus of the Galápagos Islands,Ecuador

Human activity has facilitated the introduction of a number of alien mammal species to the Galápagos Archipelago. Understanding the phylogeographic history and population genetics of invasive species on the Archipelago is an important step in predicting future spread and designing effective management strategies. In this study, we describe the invasion pathway of Rattus rattus across the Galápagos using microsatellite data, coupled with historical knowledge. Microsatellite genotypes were generated for 581 R. rattus sampled from 15 islands in the archipelago. The genetic data suggest that there are at least three genetic lineages of R. rattus present on the Galápagos Islands. The spatial distributions of these lineages correspond to the main centers of human settlement in the archipelago. There was limited admixture among these three lineages, and these finding coupled with low rates of gene flow among island populations suggests that interisland movement of R. rattus is rare. The low migration among islands recorded for the species will have a positive impact on future eradication efforts.     

Ancient DNA of the extinct lava shearwater (Puffinus olsoni) from the Canary Islands reveals incipient differentiation within the P. puffinus complex

Background

The loss of species during the Holocene was, dramatically more important on islands than on continents. Seabirds from islands are very vulnerable to human-induced alterations such as habitat destruction, hunting and exotic predators. For example, in the genus Puffinus (family Procellariidae) the extinction of at least five species has been recorded during the Holocene, two of them coming from the Canary Islands.

Methodology/Principal Findings

We used bones of the two extinct Canary shearwaters (P. olsoni and P. holeae) to obtain genetic data, for use in providing insights into the differentiation process within the genus Puffinus. Although mitochondrial DNA (mtDNA) cytochrome b sequences were successfully retrieved from four Holocene specimens of the extinct Lava shearwater (P. olsoni) from Fuerteventura (Canary Islands), the P. holeae specimens yielded no DNA. Only one haplotype was detected in P. olsoni, suggesting a low genetic diversity within this species.

Conclusions

The phylogenetic analyses based on the DNA data reveal that: (i) the “Puffinus puffinus complex”, an assemblage of species defined using osteological characteristics (P. puffinus, P. olsoni, P. mauretanicus, P. yelkouan and probably P. holeae), shows unresolved phylogenetic relationships; (ii) despite the differences in body size and proportions, P. olsoni and the extant P. puffinus are sister species. Several hypotheses can be considered to explain the incipient differentiation between P. olsoni and P. puffinus.     

Rat kappa-chain allotypes

The distribution of rat kappa-chain allotype specificities (RI-1a and 1b) was studied amongRattus rattus and a variety of other Asian rodents. No sera other than those ofRattus norvegicus showed the presence of RI-1a (DA type), whereas many cross-reacted with RI-1b (LEW-type). While manyR. rattus showedtotal cross-reactivity with RI-1b, various sera from the generaRattus, Bandicota, andTokudaia showed different levels ofpartial cross-reactivity. These results indicate that (1) anti-RI-1b reagents can detectmultiple specificities on LEW-type kappa chains, and (2) these RI-1b specificities, butnot RI-1a, are widely distributed among murid rodents, in seeming contradiction to amino acid sequence data suggesting that RI-1b is more closely related to ancestral rat kappa chains.