Evolutionary history of the little fire ant Wasmannia auropunctata before global invasion: inferring dispersal patterns,niche requirements and past and present distribution within its native range

The evolutionary history of invasive species within their native range may involve key processes that allow them to colonize new habitats. Therefore, phylogeographic studies of invasive species within their native ranges are useful to understand invasion biology in an evolutionary context. Here we integrated classical and Bayesian phylogeographic methods using mitochondrial and nuclear DNA markers with a palaeodistribution modelling approach, to infer the phylogeographic history of the invasive ant Wasmannia auropunctata across its native distribution in South America. We discuss our results in the context of the recent establishment of this mostly tropical species in the Mediterranean region. Our Bayesian phylogeographic analysis suggests that the common ancestor of the two main clades of W. auropunctata occurred in central Brazil during the Pliocene. Clade A would have differentiated northward and clade B southward, followed by a secondary contact beginning about 380 000 years ago in central South America. There were differences in the most suitable habitats among clades when considering three distinct climatic periods, suggesting that genetic differentiation was accompanied by changes in niche requirements, clade A being a tropical lineage and clade B a subtropical and temperate lineage. Only clade B reached more southern latitudes, with a colder climate than that of northern South America. This is concordant with the adaptation of this originally tropical ant species to temperate climates prior to its successful establishment in the Mediterranean region. This study highlights the usefulness of exploring the evolutionary history of invasive species within their native ranges to better understand biological invasions.     

Dispersion pattern of the medfly from its geographic centre of origin and genetic relationships of the medfly with two close relatives

We used three different approaches, a phylogenetic analysis, the principal component analysis, and Gabriel Graphs, to study the dispersion of the medfly from its geographic centre of origin. Our analysis was based on 13 enzyme polymorphisms detected electrophoretically in 15 natural populations coming from distinct geographic areas. The results indicate that (i) there is a clear-cut separation of the native African from the introduced populations, (ii) the population from Spain seems to connect the African and the Mediterranean or American populations, and (iii) the Mediterranean populations as well as the American ones comprise two distinct groups of closely related populations.We have also estimated a phylogenetic tree for 3 species of the genus Ceratitis: C. capitata, C. rosa and C. nigra, based on electrophoretic data under the assumption of a molecular clock. The results suggest that C. capitata and C. rosa are more closely related to each other than either is to C. nigra.     

Low genetic diversity of the successful invasive African clawed frog Xenopus laevis (Pipidae) in Chile

In Africa, the genus Xenopus presents cryptic species and diverse hybrids between species. It has been assumed that the invasive populations of this genus correspond to X. laevis and that they are derived from the subspecies that inhabits the Mediterranean Cape region of South Africa. In part, this is supported by the successful establishment of this species in several Mediterranean regions of the world. In Mediterranean Chile, Xenopus has invaded an area of about 21,000 km², with scarce attention to genetic aspects underlying its invasion. Using mitochondrial DNA sequences we determined that Xenopus laevis laevis from the Cape region of South Africa is the subspecies that invaded Chile. The analysis indicated that the invaders have low genetic diversity (only two haplotypes, compared to 10 in two localities of their native range), and that probably the invasion in Chile occurred only once. Landscape genetics revealed that factors such as aridity and elevation have determined the spread of the species, both from the ecological and genetic points of view. Our results show that the invasion of the African clawed frog in Chile has been successful for at least 30 years, in spite of low genetic variability, few events of introduction, low propagule pressure, and bottlenecks in the founding population.     

Evolutionary history of the European free-tailed bat,a tropical affinity species spanning across the Mediterranean Basin

The Mediterranean Basin is a global biodiversity hotspot, hosting a number of native species belonging to families that are found almost exclusively in tropical climates. Yet, whether or not these taxa were able to survive in the Mediterranean region during the Quaternary climatic oscillations remains unknown. Focusing on the European free-tailed bat (Tadarida teniotis), we aimed to (a) identify potential ancient populations and glacial refugia; (b) determine the post-glacial colonization routes across the Mediterranean; and (c) evaluate current population structure and demography. Mitochondrial and nuclear markers were used to understand T. teniotis evolutionary and demographic history. We show that T. teniotis is likely restricted to the Western Palearctic, with mitochondrial phylogeny suggesting a split between an Anatolian/Middle East clade and a European clade. Nuclear data pointed to three genetic populations, one of which is an isolated and highly differentiated group in the Canary Islands, another distributed across Iberia, Morocco, and France, and a third stretching from Italy to the east, with admixture following a pattern of isolation by distance. Evolutionary and demographic reconstruction supports a pre-Last Glacial Maximum (LGM) colonization of Italy and the Anatolian/Middle East, while the remaining populations were colonized from Italy after the Younger Dryas. We also found support for demographic expansion following the Iberian colonization. The results show that during the LGM T. teniotis persisted in Mediterranean refugia and has subsequently expanded to its current circum-Mediterranean range. Our findings raise questions regarding the physiological and ecological traits that enabled species with tropical affinities to survive in colder climates.     

Habitat niche breadth predicts invasiveness in solitary ascidians

A major focus of invasion biology is understanding the traits associated with introduction success. Most studies assess these traits in the invaded region, while only few compare nonindigenous species to the pool of potential invaders in their native region. We focused on the niche breadth hypothesis, commonly evoked but seldom tested, which states that generalist species are more likely to become introduced as they are capable of thriving under a wide set of conditions. Based on the massive introduction of tropical species into the Mediterranean via the Suez Canal (Lessepsian migration), we defined ascidians in the Red Sea as the pool of potential invaders. We constructed unique settlement plates, each representing six different niches, to assess ascidian niche breadth, and deployed them in similar habitats in the native and invaded regions. For each species found on plates, we evaluated its abundance, relative abundance across successional stages, and niche breadth, and then compared (1) species in the Red Sea known to have been introduced into the Mediterranean (Lessepsian species) and those not known from the Mediterranean (non‐Lessepsian); and (2) nonindigenous and indigenous species in the Mediterranean. Lessepsian species identified on plates in the Red Sea demonstrated wider niche breadth than non‐Lessepsian species, supporting the niche breadth hypothesis within the native region. No differences were found between Lessepsian and non‐Lessepsian species in species abundance and successional stages. In the Mediterranean, nonindigenous species numerically dominated the settlement plates. This precluded robust comparisons of niche breadth between nonindigenous and indigenous species in the invaded region. In conclusion, using Red Sea ascidians as the pool of potential invaders, we found clear evidence supporting the niche breadth hypothesis in the native region. We suggest that such patterns may often be obscured when conducting trait‐based studies in the invaded regions alone. Our findings indicate that quantifying the niche breadth of species in their native regions will improve estimates of invasiveness potential.     

MOLECULAR SYSTEMATICS OF THE GRACILARIACEAE (GRACILARIALES,RHODOPHYTA) WITH EMPHASIS ON SOUTHERN AFRICA1

Southern Africa has economically exploited populations of terete gracilarioids on the cool temperate west coast and numerous species of endemic and Indo‐Pacific tropical Gracilariaceae on the south and east coasts. Gross morphological characters have been the main means of identification, and incorrect applications have led to a number of misidentifications. In this study, small subunit rDNA and RUBISCO spacer sequences were used to determine phylogenetic relationships. Whereas rDNA sequences successfully differentiate major groups within the family as well as species belonging to the Gracilariopsis and the Curdiea/Melanthalia clade, RUBISCO spacer sequencing was required to distinguish between species of Gracilaria. The southern African gracilarioid complex (stringy, terete, elongate members of the Gracilariaceae) was resolved into three species: Gracilaria gracilis, Gracilariopsis longissima, and Gracilariopsis funicularis. South African Gracilaria protea was shown to be conspecific with tropical Indian Ocean G. corticata. Apart from G. gracilis and G. corticata, South African Gracilaria species were differentiated into a temperate‐tropical terete grouping and a temperate‐tropical flattened grouping.     

Historical biogeography of the land snail <Emphasis Type="Italic">Cornu aspersum</Emphasis>: a new scenario inferred from haplotype distribution in the Western Mediterranean basin

Background  

Despite its key location between the rest of the continent and Europe, research on the phylogeography of north African species remains very limited compared to European and North American taxa. The Mediterranean land mollusc Cornu aspersum (= Helix aspersa) is part of the few species widely sampled in north Africa for biogeographical analysis. It then provides an excellent biological model to understand phylogeographical patterns across the Mediterranean basin, and to evaluate hypotheses of population differentiation. We investigated here the phylogeography of this land snail to reassess the evolutionary scenario we previously considered for explaining its scattered distribution in the western Mediterranean, and to help to resolve the question of the direction of its range expansion (from north Africa to Europe or vice versa). By analysing simultaneously individuals from 73 sites sampled in its putative native range, the present work provides the first broad-scale screening of mitochondrial variation (cyt b and 16S rRNA genes) of C. aspersum.     

Recent Invasion of the Symbiont-Bearing Foraminifera Pararotalia into the Eastern Mediterranean Facilitated by the Ongoing Warming Trend

The eastern Mediterranean is a hotspot of biological invasions. Numerous species of Indo-pacific origin have colonized the Mediterranean in recent times, including tropical symbiont-bearing foraminifera. Among these is the species Pararotalia calcariformata. Unlike other invasive foraminifera, this species was discovered only two decades ago and is restricted to the eastern Mediterranean coast. Combining ecological, genetic and physiological observations, we attempt to explain the recent invasion of this species in the Mediterranean Sea. Using morphological and genetic data, we confirm the species attribution to P. calcariformata McCulloch 1977 and identify its symbionts as a consortium of diatom species dominated by Minutocellus polymorphus. We document photosynthetic activity of its endosymbionts using Pulse Amplitude Modulated Fluorometry and test the effects of elevated temperatures on growth rates of asexual offspring. The culturing of asexual offspring for 120 days shows a 30-day period of rapid growth followed by a period of slower growth. A subsequent 48-day temperature sensitivity experiment indicates a similar developmental pathway and high growth rate at 28°C, whereas an almost complete inhibition of growth was observed at 20°C and 35°C. This indicates that the offspring of this species may have lower tolerance to cold temperatures than what would be expected for species native to the Mediterranean. We expand this hypothesis by applying a Species Distribution Model (SDM) based on modern occurrences in the Mediterranean using three environmental variables: irradiance, turbidity and yearly minimum temperature. The model reproduces the observed restricted distribution and indicates that the range of the species will drastically expand westwards under future global change scenarios. We conclude that P. calcariformata established a population in the Levant because of the recent warming in the region. In line with observations from other groups of organisms, our results indicate that continued warming of the eastern Mediterranean will facilitate the invasion of more tropical marine taxa into the Mediterranean, disturbing local biodiversity and ecosystem structure.     

Patapius spinosus: First record of Leptopodidae (Heteroptera) from Japan

Patapius spinosus, a remarkable leptopodid bug whose body is covered with spines, was collected for the first time in Japan. This species was found in the supralittoral zone of a sandy beach in Osaka Prefecture, central Japan, in autumn 2002 and in spring 2004. Patapius spinosus is known to be distributed in the Mediterranean and North African regions, and has been introduced into North America and Chile. Its occurrence in Japan could signify either an eastern limit of its native distribution or an exotic origin.     

RADseq dataset with 90% missing data fully resolves recent radiation of Petalidium (Acanthaceae) in the ultra‐arid deserts of Namibia

Deserts, even those at tropical latitudes, often have strikingly low levels of plant diversity, particularly within genera. One remarkable exception to this pattern is the genus Petalidium (Acanthaceae), in which 37 of 40 named species occupy one of the driest environments on Earth, the Namib Desert of Namibia and neighboring Angola. To contribute to understanding this enigmatic diversity, we generated RADseq data for 47 accessions of Petalidium representing 22 species. We explored the impacts of 18 different combinations of assembly parameters in de novo assembly of the data across nine levels of missing data plus a best practice assembly using a reference Acanthaceae genome for a total of 171 sequence datasets assembled. RADseq data assembled at several thresholds of missing data, including 90% missing data, yielded phylogenetic hypotheses of Petalidium that were confidently and nearly fully resolved, which is notable given that divergence time analyses suggest a crown age for African species of 3.6–1.4 Ma. De novo assembly of our data yielded the most strongly supported and well‐resolved topologies; in contrast, reference‐based assembly performed poorly, perhaps due in part to moderate phylogenetic divergence between the reference genome, Ruellia speciosa, and the ingroup. Overall, we found that Petalidium, despite the harshness of the environment in which species occur, shows a net diversification rate (0.8–2.1 species per my) on par with those of diverse genera in tropical, Mediterranean, and alpine environments.     

A review of changes in the fish assemblages of Levantine inland and marine ecosystems following the introduction of non-native fishes

The arrival of non‐native fishes in the Levant Basin began in the late 19th century. Whereas the presence of most of the 40 non‐native freshwater fishes stem from intentional introductions, either for aquaculture or pest control, the 62 species of non‐native marine fishes arrived by natural dispersal via the Suez Canal. Of the non‐native freshwater species, five have established successful breeding populations (mosquitofish Gambusia affinis, common carp Cyprinus carpio, crucian carp Carassius carassius, swordtail Xiphophorus hellerii and rainbow trout Oncorhynchus mykiss), and seven are regularly stocked in natural habitats (thinlip mullet Liza ramada, flathead mullet Mugil cephalus, European eel Anguilla anguilla, grass carp Ctenopharyngodon idella, Asian silver carp Hypophthalmichthys molitrix, bighead carp Aristichthys nobilis, black carp Mylopharyngodon piceus). Some non‐native species appear to have out‐competed native species. Gambusia affinis may have caused the extirpation of two native cyprinid fishes from the Qishon River basin (Levant silver carp Hemigrammocapoeta nana and common garra Garra rufa) and the southern Dead Sea (endemic Sodom's garra G. ghoerensis). The opening of the Suez Canal in 1869 allowed entry into the eastern Mediterranean of Indo‐Pacific and Erythrean biota, with the latter now dominating the community structure (50–90% of fish biomass) and function (altered native food web) of the Levantine littoral and infra‐littoral zones. The process has accelerated in recent years concurrent with a warming trend of the seawater. Record numbers of newly discovered non‐native species is leading to the creation of a human‐assisted Erythrean biotic province in the eastern Mediterranean.     

Ecological impacts of invasive African olive (Olea europaea ssp. cuspidata) in Cumberland Plain Woodland,Sydney, Australia

African olive (Olea europaea ssp. cuspidata) is a small evergreen tree which has become highly invasive at a landscape scale in the western Sydney and Hunter Valley regions of New South Wales, Australia. African olive invasion results in the formation of a dense and permanent mid‐canopy in grassy woodland vegetation. We investigated the relationship between African olive and native species establishment, abundance and diversity, using field surveys and a manipulative shading experiment. There were 78% fewer native species beneath African olive canopy in the field compared to uninvaded woodland sites. The shading experiment showed that simulated dense African olive shade levels produced the lowest dry weight for the three native species studied, with simulated canopy edge light providing optimal conditions for the native shrub Bursaria spinosa and African olive. Dense African olive shade levels produced the highest mortality rate for native species; however, African olive was able to maintain an 88% survival rate under dense canopy shade. This study confirms the adaptability of African olive and its capacity to decrease native plant diversity and substantially modify native vegetation at the community level.     

The tropical history and future of the Mediterranean biota and the West African enigma

Aim Since the opening of the Suez Canal in 1869, many tropical taxa from the Indo‐West Pacific (IWP) realm have entered the Mediterranean Sea, which is experiencing rising temperatures. My aims are: (1) to compare biogeographically this tropical transformation of the Mediterranean biota with the tropical faunas of the Mediterranean and adjacent southern European and West African seas during the Late Oligocene to Pliocene interval; (2) to infer the relative contributions of the tropical eastern Atlantic and IWP to the tropical component of the marine biota in southern Europe; and (3) to understand why West Africa is not now a major source of warm‐water species. Location Southern Europe, including the Mediterranean Sea, and the coast of tropical West Africa. Methods I surveyed the literature on fossil and living shell‐bearing molluscs to infer the sources and fates of tropical subgenus‐level taxa living in southern Europe and West Africa during the Late Oligocene to Pliocene interval. Results Ninety‐four taxa disappeared from the tropical eastern Atlantic (including the Mediterranean) but persisted elsewhere in the tropics, mainly in the IWP (81 taxa, 86%) and to a lesser extent in tropical America (36 taxa, 38%). Nine taxa inferred to have arrived in the tropical eastern Atlantic from the west after the Pliocene did not enter the Mediterranean. The modern West African fauna is today isolated from that of other parts of the marine tropics. Main conclusions Taxa now entering the Mediterranean through the Suez Canal are re‐establishing a link with the IWP that last existed 16 million years ago. This IWP element, which evolved under oligotrophic conditions and under a regime of intense anti‐predatory selection, will continue to expand in the increasingly warm and increasingly oligotrophic Mediterranean. The IWP source fauna contrasts with the tropical West African biota, which evolved under productive conditions and in a regime of less anti‐predatory specialization. Until now, the post‐Pliocene West African source area has been isolated from the Mediterranean by cold upwelling. If further warming should reduce this barrier, as occurred during the productive and warm Early Pliocene, the Mediterranean could become a meeting place for two tropical faunas of contrasting source conditions.     

mtDNA sequences of Sphyraena viridensis (Perciformes: Sphyraenidae) from Italy: insights into historical events and the phylogeny of the genus

The yellowmouth barracuda, Sphyraena viridensis, is a Mediterranean native species whose exact distribution is uncertain due to a long‐term taxonomic confusion with Sphyraena sphyraena. Records of this species in the Mediterranean Sea have recently increased, and a northwards expansion of its distribution has been suggested. Three mtDNA regions, namely cytochrome oxidase I, cytochrome b and the control region, were analysed in S. viridensis samples from Italian coastal regions to provide molecular markers useful in species identification, in phylogenetic analysis and in detecting the distribution of genetic variability of the yellowmouth barracuda in this area. The data clearly distinguish S. viridensis from S. sphyraena and the other four (one native and three Lessepsian) Mediterranean Sphyraena species and identify two clearly distinct lineages that diverged during the Pleistocene but are currently panmictic in the investigated area. Both lineages retain signatures of historical population expansion. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 635–641.     

A tale of the nearly tail‐less: the effects of Plio‐Pleistocene climate change on the diversification of the African avian genus Sylvietta

Sylvietta is a broadly distributed group of African species inhabiting a wide range of habitats and presents an interesting opportunity to investigate the historic mechanisms that have impacted the biogeography of African avian species. We collected sequence data from 50 individuals and used model‐based phylogenetic methods, molecular divergence estimates and ancestral area estimates to construct a time‐calibrated phylogeny and estimation of biogeographic history. We estimate a southern African origin for Sylvietta, with an initial divergence splitting the genus into two clades. The first consists of arid‐adapted species, with a southern African origin and subsequent diversification north into Ethiopia–Somalia. The second clade is estimated as having a Congolian forest origin with an eastward pattern of colonization and diversification as a result of Plio‐Pleistocene forest dynamics. Additionally, two members of the genus Sylvietta display interesting patterns of intraspecific diversification. Sylvietta rufescens is an arid‐adapted species inhabiting southern Africa, and we recover two subclades with a divergence dating to the Pleistocene, a unique pattern for avian species which may be explained via isolation in arid habitat fragments in the early Pleistocene. Second, Sylvietta virens, a species endemic to Afro‐tropical forests, is recovered with geographically structured genetic diversification across its broad range, an interesting result given that recent investigations of several avian forest species have found similar and substantial geographically structured genetic diversity relating to Plio‐Pleistocene forest fragmentation. Overall, Plio‐Pleistocene habitat cycling played a significant role in driving diversification in Sylvietta, and this investigation highlights the substantial impact of climate‐driven habitat dynamics on the history of sub‐Saharan species.     

Tectonics,climate and the diversification of the tropical African terrestrial flora and fauna

Tropical Africa is home to an astonishing biodiversity occurring in a variety of ecosystems. Past climatic change and geological events have impacted the evolution and diversification of this biodiversity. During the last two decades, around 90 dated molecular phylogenies of different clades across animals and plants have been published leading to an increased understanding of the diversification and speciation processes generating tropical African biodiversity. In parallel, extended geological and palaeoclimatic records together with detailed numerical simulations have refined our understanding of past geological and climatic changes in Africa. To date, these important advances have not been reviewed within a common framework. Here, we critically review and synthesize African climate, tectonics and terrestrial biodiversity evolution throughout the Cenozoic to the mid-Pleistocene, drawing on recent advances in Earth and life sciences. We first review six major geo-climatic periods defining tropical African biodiversity diversification by synthesizing 89 dated molecular phylogeny studies. Two major geo-climatic factors impacting the diversification of the sub-Saharan biota are highlighted. First, Africa underwent numerous climatic fluctuations at ancient and more recent timescales, with tectonic, greenhouse gas, and orbital forcing stimulating diversification. Second, increased aridification since the Late Eocene led to important extinction events, but also provided unique diversification opportunities shaping the current tropical African biodiversity landscape. We then review diversification studies of tropical terrestrial animal and plant clades and discuss three major models of speciation: (i) geographic speciation via vicariance (allopatry); (ii) ecological speciation impacted by climate and geological changes, and (iii) genomic speciation via genome duplication. Geographic speciation has been the most widely documented to date and is a common speciation model across tropical Africa. We conclude with four important challenges faced by tropical African biodiversity research: (i) to increase knowledge by gathering basic and fundamental biodiversity information; (ii) to improve modelling of African geophysical evolution throughout the Cenozoic via better constraints and downscaling approaches; (iii) to increase the precision of phylogenetic reconstruction and molecular dating of tropical African clades by using next generation sequencing approaches together with better fossil calibrations; (iv) finally, as done here, to integrate data better from Earth and life sciences by focusing on the interdisciplinary study of the evolution of tropical African biodiversity in a wider geodiversity context.     

Flower development and reproductive continuity in MediterraneanRuscus aculeatus L. (Liliaceae)

Summary Ruscus aculeatus is a subandroecious species widespread in Mediterranean environments. Our studies on floral differentiation show that the unisexual flowers pass through a stage in which both androecial and gynoecial primordia are initiated. The hypothesis that dioecy has arisen secondarily from hermaphroditism is discussed. Data on flowering phenology of Mediterranean populations ofR. aculeatus showed it has a long flowering season (about 7 months). This phenology is similar to that of the Mediterranean species originated before the development of the Mediterranean climate. Members of the family Ruscaceae where present in Laurasia during the tropical Tertiary and the present study on flower and fruit morphology and reproductive phenology reveals forR. aculeatus a list of characters generally reported for tropical species. Small and greenish flowers, fleshy fruits, few large seeds, and resprouting capacity, together with long flowering season and continuous availability of ripe fruits, occur all together inR. aculeatus. It constitutes a tropical reproductive syndrome which might have survived the climatic fluctuations of the Quaternary, the establishment of the Mediterranean climate, and the present anthropogenic disturbs.     

Distribution and biogeography of the Paradiaptominae (Copepoda: Calanoida: Diaptomidae)

The Paradiaptominae, a subfamily of the large Diaptomidae family, is a small group of freshwater calanoids of which 24 species in four genera have been described. They are endemic to Africa with the exception of four species, Metadiaptomus asiaticus (Asia and Mongolia), M. gauthieri (Africa and Madagascar), Paradiaptomus greeni (India and Sri Lanka) and Neolovenula alluaudi (Canary Islands, North Africa, Baltic and Mediterranean countries). The North African species Metadiaptomus chevreuxi extends its range into Iraq and Iran. The Paradiaptominae are widely distributed in semi-permanent water bodies in the drier areas of Africa although most individual species have a restricted distribution. Only a few species have the ability to colonise permanent waters and these species are usually located at higher altitudes. The Paradiaptominae have not been recorded from the wet equatorial lowland areas of Africa. It seems likely that they are Gondwanian in origin as evidenced by their distribution (Africa, Madagascar, India and Sri Lanka). Neolovenula alluaudi (Mediterranean) shares morphological characters tenuously linked to the other species in the group and may either have a different origin or have branched off at an early stage in evolution of the group.     

Mammal distributions in the western Mediterranean: the role of human intervention

The natural distribution of the 17 non-flying mammal species occurring wild in both the Maghreb (north-west Africa) and Iberia (south-west Europe) is considered. It is concluded that only four species – Red Fox Vulpes vulpes, Wild Boar Sus scrofa, Wild Cat Felis silvestris and Otter Lutra lutra – are native to both regions, while another three – Red Deer Cervus elaphus, Brown Bear Ursus arctos and Aurochs Bos primigenius – were native to North Africa until the mid-Holocene but have probably died out naturally. Algerian Hedgehog Atelerix algirus, Barbary Ape Macaca sylvanus, Genet Genetta genetta and Egyptian Mongoose Herpestes ichneumon are widely accepted as introductions to Europe from North Africa. The remaining six species, and Red Deer now found in Africa, were also probably introduced – Rabbit Oryctolagus cuniculus, Weasel Mustela nivalis, Wood Mouse Apodemus sylvaticus and Lesser White-toothed Shrew Crocidura russula from Europe to Africa; Algerian Mouse Mus spretus from Africa to Europe; Savi’s Pygmy Shrew Suncus etruscus perhaps from the eastern Mediterranean to both Iberia and the Maghreb. There are two Maghrebi species which, although not found in Europe, are more closely related to Palaearctic than to Afrotropical species: Garden Dormouse Eliomys melanurus, probably native to north-west Africa, although possible augmentation of the natural population cannot be ruled out, and Whitaker’s Shrew Crocidura whitakeri, a North African endemic. Removal of so many species of European provenance from the list of mammals native to north-west Africa should not be considered to weaken its position as part of the Palaearctic zoogeographical region. Bats and other, non-mammalian, taxa illustrate the clear faunal relationship between the Maghreb and south-west Europe, whilst emphasizing its distinction from subsaharan Africa.