rbcL sequences reveal multiple cryptic introductions of the Japanese red alga Polysiphonia harveyi

In Europe, the last 20 years have seen a spectacular increase in accidental introductions of marine species, but it has recently been suggested that both the actual number of invaders and their impacts have been seriously underestimated because of the prevalence of sibling species in marine habitats. The red alga Polysiphonia harveyi is regarded as an alien in the British Isles and Atlantic Europe, having appeared in various locations there during the past 170 years. Similar or conspecific populations are known from Atlantic North America and Japan. To choose between three competing hypotheses concerning the origin of P. harveyi in Europe, we employed rbcL sequence analysis in conjunction with karyological and interbreeding data for samples and isolates of P. harveyi and various congeners from the Pacific and North Atlantic Oceans. All cultured isolates of P. harveyi were completely interfertile, and there was no evidence of polyploidy or aneuploidy. Thus, this biological species is both morphologically and genetically variable: intraspecific rbcL divergences of up to 2.1% are high even for red algae. Seven rbcL haplotypes were identified. The four most divergent haplotypes were observed in Japanese samples from Hokkaido and south-central Honshu, which are linked by hypothetical 'missing' haplotypes that may be located in northern Honshu. These data are consistent with Japan being the centre of diversity and origin for P. harveyi. Two non-Japanese lineages were linked to Hokkaido and Honshu, respectively. A single haplotype was found in all North Atlantic and Mediterranean accessions, except for North Carolina, where the haplotype found was the same as that invading in New Zealand and California. The introduction of P. harveyi into New Zealand has gone unnoticed because P. strictissima is a morphologically indistinguishable native sibling species. The sequence divergence between them is 4-5%, greater than between some morphologically distinct red algal species. Two different types of cryptic invasions of P. harveyi have therefore occurred. In addition to its introduction as a cryptic sibling species in New Zealand, P. harveyi has been introduced at least twice into the North Atlantic from presumed different source populations. These two introductions are genetically and probably also physiologically divergent but completely interfertile.     

AN ATLANTIC POPULATION OF PIKEA CALIFORNICA (DUMONTIACEAE,RHODOPHYTA)1

The marine red alga Pikea californica Harvey, previously known only from the east coast of Japan and the west coast of North America, was found in 1983 to be abundant in the surge zone throughout the Isles of Scilly archipelago, off SW England. Examination of herbarium material showed that the population was present in 1967. All plants observed in July 1983 and September 1984 were sterile, but reproductive male and female plants were collected in November 1983. The habitat and phenology of P. californica in the Isles of Scilly closely resemble those of Pacific populations. English plants are vegetatively and reproductively identical to the type and other California specimens. They can easily be distinguished from Sphaerococcus coronopifolius Stackhouse (Sphaerococcaceae, Gigartinales), the most similar species in the flora of the British Isles, by the presence in transverse sections of several lacunae, representing axial and whorl-branchlet filaments, in contrast to the single axial filament of S. coronopifolius. It is possible that the Isles of Scilly population of P. californica is a relict of a species once more widespread, but it seems more likely that it is another example of the well-established pattern of marine introductions into the British Isles from the North Pacific. The range of temperature regimes occupied by P. Californica in the Pacific suggests that the species could become more widespread in Europe, but at present it appears to be confined to the Isles of Scilly.     

AN ATLANTIC POPULATION OF PIKEA CALIFORNICA (DUMONTIACEAE,RHODOPHYTA)1

The marine red alga Pikea californica Harvey, previously known only from the east coast of Japan and the west coast of North America, was found in 1983 to be abundant in the surge zone throughout the Isles of Scilly archipelago, off SW England. Examination of herbarium material showed that the population was present in 1967. All plants observed in July 1983 and September 1984 were sterile, but reproductive male and female plants were collected in November 1983. The habitat and phenology of P. californica in the Isles of Scilly closely resemble those of Paczfic populations. English plants are vegetatively and reproductively identical to the type and other California specimens. They can easily be distinguished from Sphaerococcus coronopifolius Stackhouse (Sphaerococcaceae, Gigartinales), the most similar species in the flora of the British Isles, by the presence in transverse section of several lacunae, representing axial and whorlbranchletjilaments, in contrast to the single axial filament of S. coronopifolius. It is possible that the Isles of Scilly population of P. californica is a relict of a species once more widespread, but it seems more likely that it is another example of the well-established pattern of marine introductions into the British Isles from the North Paczfic. The range of temperature regimes occupied by P. californica in the Pacific suggests that the species could become more widespread in Europe, but at present it appears to be confined to the Isles of Scilly.     

GLOBAL SURVEY OF MITOCHONDRIAL DNA SEQUENCES IN THE THREESPINE STICKLEBACK: EVIDENCE FOR RECENT MIGRATIONS

Phylogenetic analyses of mitochondrial DNA (mtDNA) sequences were used to assess the matriarchal genetic structure of the threespine stickleback, Gasterosteus aculeatus. A 747 base-pair (bp) fragment of the cytochrome b was sequenced from 36 individuals collected from 25 localities in Europe, North America, and Japan. Two major divergent clades were revealed: one widespread in Japan but with representatives in some Alaskan and British Columbian lakes and the other common in Europe and North America. A simple diagnostic test using the polymerase chain reaction (PCR) and a restriction enzyme was used to assay additional individuals, confirming the absence of the Japanese clade in the Atlantic basin. Geographic distribution of mtDNA variation suggests (1) a recent origin of the Atlantic populations, and (2) support for previous hypotheses about the existence of Pleistocene refugia for freshwater fishes in Alaska and British Columbia. Silent substitution rates were used to date the colonization of the Atlantic at 90,000 to 260,000 yr before present, which conflicts with earlier dates implied by the fossil record. The recent replacement of Atlantic mitochondrial lineages suggested by our data may be explained by severe reduction or extinction of northern Atlantic populations during the Pleistocene, followed by a recent reinvasion from the Pacific. With a global perspective of the distribution of genetic variation as a framework, meaningful comparisons at a smaller geographical scale will now be possible.     

A new xanthone-containing Micarea from Northwest Europe and the Pacific Northwest of North America

Abstract:Micarea xanthonica sp. nov. is described from the western British Isles, westernmost Norway and the Pacific Northwest of North America. It belongs to the M. prasina group, but differs from related species in possessing xanthones in the thallus.     

From Europe to America: pliocene to recent trans-atlantic expansion of cold-water north atlantic molluscs

Data on the geographical distribution, phylogeny and fossil record of cool-temperate North Atlantic shell-bearing molluscs that live in waters shallower than 100 m depth belong to two biogeographic provinces, one in eastern North America north of Cape Cod, the other in northern Europe. Amphi-Atlantic species, which are found in both provinces, comprise 30.8% of the 402 species in the northeastern Atlantic and 47.3% of the 262 species in the northwestern Atlantic. Some 54.8% of these amphi-Atlantic species have phylogenetic origins in the North Pacific. Comparisons among fossil Atlantic faunas show that amphi-Atlantic distributions became established in the Middle Pliocene (about 3.5 million years ago), and that all represent westward expansions of European taxa to North America. No American taxa spread eastward to Europe without human assistance. These results are in accord with previous phylogeographic studies among populations within several amphi-Atlantic species. Explanations for the unidirectional expansion of species across the Atlantic remain uncertain, but may include smaller size and greater prior extinction of the North American as compared to the European fauna and biased transport mechanisms. Destruction of the European source fauna may jeopardize faunas on both sides of the Atlantic.     

PORPHYRA SUBORBICULATA, PORPHYRA CAROLINENSIS AND PORPHYRA LILLIPUTIANA - THREE NAMES FOR ONE SMALL PORPHYRA

The genus Porphyra is ancient, successful, and morphologically simple. Its members provide a particular challenge to systematists who must decide whether shared features are a result of homoplasy, or reflect recent common ancestry. Three species of diminutive Porphyra with widespread geographic origins share many common morphological features: P suborbiculata Kjellm. has been reported from the west Pacific and Indian Oceans, P. carolinensis Coll et J. Cox from the west Atlantic, and P lilliputiana W. A. Nelson, G. A.Knight et M. W. Hawkes from New Zealand. Comparison of 18S rDNA sequence data from small Porphyra thalli from Mexico, Japan, New Zealand, Australia and Connecticut indicate that these three taxa in fact belong to one cosmopolitan species, which is distributed over three oceans and two hemispheres. Analysis of sequence data from introns present in the 18S rDNA and from the ITS region suggest that this distribution may be linked to human activity, and raises the question of to which geographic locality this entity is truly endemic?     

Patterns of genetic diversity of the cryptogenic red alga Polysiphonia morrowii (Ceramiales,Rhodophyta) suggest multiple origins of the Atlantic populations

The red alga Polysiphonia morrowii, native to the North Pacific (Northeast Asia), has recently been reported worldwide. To determine the origin of the French and Argentine populations of this introduced species, we compared samples from these two areas with samples collected in Korea and at Hakodate, Japan, the type locality of the species. Combined analyses of chloroplastic (rbcL) and mitochondrial (cox1) DNA revealed that the French and Argentine populations are closely related and differ substantially from the Korean and Japanese populations. The genetic structure of P. morrowii populations from South Atlantic and North Atlantic, which showed high haplotype diversity compared with populations from the North Pacific, suggested the occurrence of multiple introduction events from areas outside of the so‐called native regions. Although similar, the French and Argentine populations are not genetically identical. Thus, the genetic structure of these two introduced areas may have been modified by cryptic and recurrent introduction events directly from Asia or from other introduced areas that act as introduction relays. In addition, the large number of private cytoplasmic types identified in the two introduced regions strongly suggests that local populations of P. morrowii existed before the recent detection of these invasions. Our results suggest that the most likely scenario is that the source population(s) of the French and Argentine populations was not located only in the North Pacific and/or that P. morrowii is a cryptogenic species.     

Mitochondrial DNA reveals multiple Northern Hemisphere introductions of Caprella mutica (Crustacea, Amphipoda)

Caprella mutica (Crustacea, Amphipoda) has been widely introduced to non-native regions in the last 40 years. Its native habitat is sub-boreal northeast Asia, but in the Northern Hemisphere, it is now found on both coasts of North America, and North Atlantic coastlines of Europe. Direct sequencing of mitochondrial DNA (cytochrome c oxidase subunit I gene) was used to compare genetic variation in native and non-native populations of C. mutica . These data were used to investigate the invasion history of C. mutica and to test potential source populations in Japan. High diversity (31 haplotypes from 49 individuals), but no phylogeographical structure, was identified in four populations in the putative native range. In contrast, non-native populations showed reduced genetic diversity (7 haplotypes from 249 individuals) and informative phylogeographical structure. Grouping of C. mutica populations into native, east Pacific, and Atlantic groups explained the most among-region variation (59%). This indicates independent introduction pathways for C. mutica to the Pacific and Atlantic coasts of North America. Two dominant haplotypes were identified in eastern and western Atlantic coastal populations, indicating several dispersal routes within the Atlantic. The analysis indicated that several introductions from multiple sources were likely to be responsible for the observed global distribution of C. mutica , but the pathways were least well defined among the Atlantic populations. The four sampled populations of C. mutica in Japan could not be identified as the direct source of the non-native populations examined in this study. The high diversity within the Japan populations indicates that the native range needs to be assessed at a far greater scale, both within and among populations, to accurately assess the source of the global spread of C. mutica .     

The distribution and reproduction of Pterosiphonia complanata

The geographical distribution of the red alga Pterosiphonia complanata (Clem.) Falkenb. is described. New records confirm that it is restricted to the warmer waters of the North Atlantic and extend its known distribution in the British Isles where it is at the northern limits of its range. The apparent scarcity and restricted distribution of the species in the British Isles and the absence of sexual plants throughout its range are discussed in relation to the habitat and possible life history of the species.     

Evidence for the introduction of the Asian red alga Neosiphonia japonica and its introgression with Neosiphonia harveyi (Ceramiales,Rhodophyta) in the Northwest Atlantic

There is currently conflict in the literature on the taxonomic status of the reportedly cosmopolitan species Neosiphonia harveyi, a common red alga along the coast of Atlantic Canada and New England, USA. Neosiphonia harveyi sensu lato was assessed using three molecular markers: COI‐5P, ITS and rbcL. All three markers clearly delimited three genetic species groups within N. harveyi sensu lato in this region, which we identified as N. harveyi, N. japonica and Polysiphonia akkeshiensis (here resurrected from synonymy with N. japonica). Although Neosiphonia harveyi is considered by some authors to be introduced to the Atlantic from the western Pacific, it was only confirmed from the North Atlantic suggesting it is native to this area. In contrast, Neosiphonia japonica was collected from only two sites in Rhode Island, USA, as well as from its reported native range in Asia (South Korea), which when combined with data in GenBank indicates that this species was introduced to the Northwest Atlantic. The GenBank data further indicate that N. japonica was also introduced to North Carolina, Spain, Australia and New Zealand. Despite the fact that all three markers clearly delimited N. harveyi and N. japonica as distinct genetic species groups, the ITS sequences for some N. harveyi individuals displayed mixed patterns and additivity indicating introgression of nuclear DNA from N. japonica into N. harveyi in the Northwest Atlantic. Introgression of DNA from an introduced species to a native species (i.e. ‘genetic pollution’) is one of the possible consequences of species introductions, and we believe this is the first documented evidence for this phenomenon in red algae.     

Patterns of mitochondrial haplotype diversity in the invasive pest Epiphyas postvittana (Lepidoptera: Tortricidae)

The light brown apple moth, Epiphyas postvittana (Walker) (Lepidoptera: Tortricidae), is a horticultural pest of Australia and New Zealand that has more recently invaded Hawaii, Europe, and California. A 2,216-bp region of the mitochondrial genome containing the cytochrome oxidase I and II genes was sequenced from 752 individuals. Haplotype network analyses revealed a major split between a predominantly Western Australian clade and all other samples, suggestive of either a deep genetic divergence or a cryptic species. Nucleotide and haplotype diversity were highest in the country of origin, Australia, and in New Zealand populations, with evidence of haplotype sharing between New Zealand and Tasmania. Nucleotide and haplotype diversity were higher in California than within the British Isles or Hawaii. From the total of 96 haplotypes, seven were found in California, of which four were private. Within California, there have been at least two introductions; based on genetic diversity we were unable to assign a likely source for a single moth found and eradicated in Los Angeles in 2007; however, our data suggest it is unlikely that Hawaii and the British Isles are sources of the major E. postvittana population found throughout the rest of the state since 2006.     

Indian migrant butterflies displaced to Arabia by monsoon storm 'Aurora' in August 1983

Although many insect species are now thought to travel hundreds and even thousands of kilometres on the wind (Pedgley, 1982) only a few are butterflies. Three species for which there is reasonably convincing but circumstantial evidence are Danaus plexippus L. from the U.S.A. to the British Isles (Hurst, 1969), Hypolimnas bolina nerina (F.) from Australia to New Zealand (Tomlinson, 1973) and Nymphalis antiopa L. from northern Europe to the British Isles (Chalmers-Hunt, 1977). In this paper we describe the sudden appearance of Indian butterflies in Arabia and provide evidence that they were windborne across the Arabian Sea.     

A taxonomic,biogeographical and ecological overview of invasive woody plants

Abstract. Invasive plants are considered to be a major threat to the earth's biodiversity, but have not been sufficiently investigated. To address this problem a relational database on invasive woody plants has been set up. It is based on 2045 bibliographical references and contains records on 653 species representing 110 families. The families with the largest number of invasive species are: Rosaceae, Mimosaceae, Papilionaceae and Pinaceae. Out of 1060 recorded invasive events an equal number are reported from continents and oceanic islands. The highest number of invasive woody species are recorded from Europe, followed by the Pacific islands, North America, New Zealand, Australia, Indian Ocean islands and southern Africa. Included in these regions are areas which have fewer highly invasive species, e.g. islands on continental shelves, such as the British Isles. Although most invasions occur in disturbed habitats, most natural communities are susceptible to woody plant invasions. Data on species attributes are only available for a minority of species but indicate that invasive woody plant species may be either insect or wind-pollinated, have a wide array of fruit types, fruit and seed sizes, number of seeds per fruit and dispersal agents. The relative frequency of several attributes varies with the degree of invasiveness. Thus it is not yet possible to determine which set of attributes favours invasiveness and therefore it is difficult to make predictions.     

THE PELAGIC DISTRIBUTION OF THE SOOTY SHEARWATER PROCELLARIA GRISEA

The breeding distribution of the Sooty Shearwater and the dates of its breeding season are briefly reviewed.
Records from the Southern Ocean are summarised. It is concluded that they are accounted for by non-breeding birds feeding off the ice-edge in the Australian sector of the Antarctic during the latter part of the breeding season.
The great majority of Sooty Shearwaters spend the southern winter in the Pacific Ocean. Birds from the New Zealand colonies are probably mostly found on the feeding grounds off Japan: some may make a "circular" migration around the Pacific. Birds from the Cape Horn colonies spend the southern winter in the rich areas off the western coast of North America.
Some Sooty Shearwaters spend the southern winter off the South African coasts. A number of non-breeding birds remain there throughout the summer.
The Sooty Shearwaters "wintering" in the North Atlantic are probably numbered in tens of thousands only. The distribution of records is reviewed month by month and the records from Atlantic transects are summarised. The migrations in the North Atlantic are discussed in relation to prevailing meteorological and oceanographic conditions, and food supply. It is concluded that most Sooty Shearwaters in the North Atlantic make a "circular" migration, crossing the North Atlantic from west to east in June and July, and re-crossing about September further south in the northeast trades.     

Potential transmission of West Nile virus in the British Isles: an ecological review of candidate mosquito bridge vectors

West Nile virus (WNV) transmitted by mosquitoes (Diptera: Culicidae) infects various vertebrates, being pathogenic for birds, horses and humans. After its discovery in tropical Africa, sporadic outbreaks of WNV occurred during recent decades in Eurasia, but not the British Isles. WNV reached New York in 1999 and spread to California by 2003, causing widespread outbreaks of West Nile encephalitis across North America, transmitted by many species of mosquitoes, mainly Culex spp. The periodic reappearance of WNV in parts of continental Europe (from southern France to Romania) gives rise to concern over the possibility of WNV invading the British Isles. The British Isles have about 30 endemic mosquito species, several with seasonal abundance and other eco-behavioural characteristics predisposing them to serve as potential WNV bridge vectors from birds to humans. These include: the predominantly ornithophilic Culex pipiens L. and its anthropophilic biotype molestus Forskal; tree-hole adapted Anopheles plumbeus Stephens; saltmarsh-adapted Ochlerotatus caspius Pallas, Oc. detritus Haliday and Oc. dorsalis (Meigen); Coquillettidia richiardii Ficalbi, Culiseta annulata Schrank and Cs. morsitans (Theobald) from vegetated freshwater pools; Aedes cinereus Meigen, Oc. cantans Meigen and Oc. punctor Kirby from seasonal woodland pools. Those underlined have been found carrying WNV in other countries (12 species), including the rarer British species Aedes vexans (Meigen), Culex europaeus Ramos et al., Cx. modestus Ficalbi and Oc. sticticus (Meigen) as well as the Anopheles maculipennis Meigen complex (mainly An. atroparvus van Thiel and An. messeae Falleroni in Britain). Those implicated as key vectors of WNV in Europe are printed bold (four species). So far there is no proof of any arbovirus transmission by mosquitoes in the British Isles, although antibodies to Sindbis, Tahyna, Usutu and West Nile viruses have been detected in British birds. Neighbouring European countries have enzootic WNV and human infections transmitted by mosquito species that are present in the British Isles. However, except for localized urban infestations of Cx. pipiens biotype molestus that can be readily eliminated, there appear to be few situations in the British Isles where humans and livestock are exposed to sustained risks of exposure to potential WNV vectors. Monitoring of mosquitoes and arbovirus surveillance are required to guard the British Isles against WNV outbreaks and introduction of more anthropophilic mosquitoes such as Stegomyia albopicta (Skuse) and Ochlerotatus japonicus (Theobald) that have recently invaded Europe, since they transmit arboviruses elsewhere.     

Morphological and molecular characterisation of Puccinia lagenophorae, now present in central North America

In June 2004, a rust fungus not previously reported for Oklahoma was found occurring naturally on the weed, common groundsel, Senecio vulgaris, in pots in a commercial container nursery in northeastern Oklahoma. Host symptoms and morphology of teliospores and aeciospores of the fungus were consistent with those of Puccinia lagenophorae, a recent introduction into North America that has, as yet, been reported only on the East and West Coasts of the USA. This is the first report of the rust in central regions of North America. The rust is believed to be native to Australia and New Zealand and subsequently reported in most continents on numerous species and genera of the Asteraceae. Some authors in Europe consider the rust on Bellis as different from the one on Senecio, naming it Puccinia distincta. Our ITS1‐5.8S‐ITS2 ribosomal DNA sequence data, however, show homology with P. distincta sequences from Europe, indicating there is only one morphologically‐variable polyphagous species. Presumably, the rust was introduced into Oklahoma on infected plants from the West Coast of the USA, the source of most plant material in the commercial nursery where it was found. The rust is potentially important on several ornamental Asteraceae in North America as it has become in Europe, where it spreads rapidly throughout that continent following its introduction there in the early 1960s.     

Windborne displacements of Desert Locusts from Africa to the Caribbean and South America

The Desert Locust is a major pest of agriculture in Africa, the Middle East and South-West Asia and swarms are known to make downwind flights over hundreds and thousands of kilometres between seasonal breeding areas. At the end of summer in 1988, swarms of locusts were moving north and south along the western margins of North Africa and in October and November, swarms crossed the Atlantic Ocean and invaded the Caribbean and neighbouring parts of South America for the first recorded time. Because of the extent of the migration and the evolutionary significance of linkages between Old and New World species of locusts, the weather associated with the migrations was studied and trajectory analysis was used to identify the source areas and estimate the flight times. Locusts were moving offshore from western North Africa throughout the autumn and on three occasions migrated west of 40° W with easterly Trade winds. Two trans-Atlantic crossings coincided with the passage of easterly waves. Over 100 trajectories were constructed at 950 and 850 hPa and within the time limit used ( 144-h), 28% successfully linked source and receptor areas. Minimum trajectory duration was 93-h, which is one-and-a-half times longer than the previously longest flight duration, derived for a similar migration to the British Isles in 1954. Upwind trajectories from the arrival areas, identified sources between 27 and 6° N in Africa, with most end-points located in Mauritania, Senegal, Gambia and Guinea-Bissau. Interspersed with the Atlantic crossings were a northward movement of locusts and an incursion of Saharan dust into Europe within the circulations of frontal depressions. While offshore migrations from northern Africa are common in autumn, the immigrants in the Caribbean and South America were probably at the extreme limits of flight endurance for the species. The results tend to confirm earlier hypotheses that New World species of locusts may have evolved from ancestral migrants from Africa.     

GENETIC DATA HINT AT A COMMON DONOR REGION FOR INVASIVE ATLANTIC AND PACIFIC POPULATIONS OF GRACILARIA VERMICULOPHYLLA (GRACILARIALES,RHODOPHYTA)1

Gracilaria vermiculophylla (Ohmi) Papenf., an agar‐producing red alga introduced from northeast Asia to Europe and North America, is often highly abundant in invaded areas. To assay its genetic diversity and identify the putative source of invasive populations, we analyzed the mitochondrial cytochrome c oxidase subunit I (cox1) gene from 312 individuals of G. vermiculophylla collected in 37 native and 32 introduced locations. A total of 19 haplotypes were detected: 17 in northeast Asia and three in Europe and eastern and western North America, with only one shared among all regions. The shared haplotype was present in all introduced populations and in ～99% of individuals in the introduced areas. This haplotype was also found at three native locations in east Korea, west Japan, and eastern Russia. Both haplotype and nucleotide diversities were extremely low in Europe and North America compared to northeast Asia. Our study indicates that the East Sea/Sea of Japan is a likely donor region of the invasive populations of G. vermiculophylla in the east and west Atlantic and the east Pacific.     

Molecular phylogeny of the Sphaerophorus globosus species complex

The Sphaerophorus globosus complex (Lecanorales, lichenized Ascomycota) shows a large morphological variation, and three relatively distinct morphotypes can be distinguished in parts of the distribution area. Here, we utilize a multigene‐based maximum‐parsimony approach (nITS+ LSU rDNA, mtSSU rDNA, β‐tubulin, and actin) to investigate whether these morphotypes constitute distinct species. The results show that there are at least two well‐supported monophyletic groups that we interpret as phylogenetic species within the S. globosus complex. These species do not completely correspond to the predefined morphotypes. One group, an apparently undescribed species, contains noncoralloid specimens from the North American Pacific Northwest. The other group, S. globosus, consists of two well‐supported monophyletic groups: one contains coralloid epiphytic specimens from the North American Pacific Northwest that are morphologically indistinguishable from epiphytic specimens from Europe and are presently interpreted as belonging to the same species and the other is morphologically variable and contains terrestrial specimens from Europe, North America, and southernmost South America and coralloid epiphytic and epilithic specimens from Europe. The results suggest that the population in southernmost South America originated by long‐distance dispersal from arctic populations in the Northern Hemisphere.